A quantitative analysis of sources for summertime phytoplankton variability over 18 years in the South Shetland Islands (Antarctica) region

Eighteen years of summertime hydrographic and chlorophyll-a (Chl-a) data (～2700 stations) from the South Shetland Islands (Antarctica) region show that a “bell-shaped” (unimodal) distribution of phytoplankton biomass results annually when plotted against the inshore to offshore gradient in surface salinity. The maximum for this unimodal Chl-a distribution corresponds with a shallow upper mixed layer (UML) in iron-rich waters that occurs at salinities ～34. Methods of gradient analysis are used to distinguish sources of variability for bloom development among years. The control of phytoplankton biomass is resolved across the salinity gradient that separates the co-limiting conditions of deep UML depths and low-iron concentrations as opposing end-members. Chlorophyll-fluorescence yield data (a proxy for Fe-stress) showed that at salinities ～34, phytoplankton biomass was unlikely to be limited by Fe. Instead, blooming at salinities ～34 (1.3±1 mg Chl-a m^?3) co-varied with shallow UML depths (41±19 m) that occurred as a function of higher UML temperature (1.5±0.5 °C) among years, and is evidence that atmospheric climate variability impacts summertime phytoplankton biomass and production in this Southern Ocean seascape.     

Microzooplankton grazing impact in the Western Arctic Ocean

Microzooplankton grazing impact on phytoplankton was assessed using the Landry–Hassett dilution technique in the Western Arctic Ocean during spring and summer 2002 and 2004. Forty experiments were completed in a region encompassing productive shelf regions of the Chukchi Sea, mesotrophic slope regions of the Beaufort Sea off the North Slope of Alaska, and oligotrophic deep-water sites in the Canada Basin. A variety of conditions were encountered, from heavy sea-ice cover during both spring cruises, moderate sea-ice cover during summer of 2002, and light to no sea ice during summer of 2004, with a concomitant range of trophic conditions, from low chlorophyll-a (Chl-a; <0.5 μg L⁻¹) during heavy ice cover in spring and in the open basin, to late spring and summer shelf and slope open-water diatom blooms with Chl-a >5 μg L⁻¹. The microzooplankton community was dominated by large naked ciliates and heterotrophic gymnodinoid dinoflagellates. Significant, but low, rates of microzooplankton herbivory were found in half of the experiments. The maximum grazing rate was 0.16 d⁻¹ and average grazing rate, including experiments with no significant grazing, was 0.04±0.06 d⁻¹. Phytoplankton intrinsic growth rates varied from the highest values of about 0.4 d⁻¹ to the lowest values of zero to slightly negative growth, on average 0.16±0.15 d⁻¹. Light limitation in spring and post-bloom senescence during summer were likely explanations of observed low phytoplankton growth rates. Microzooplankton grazing consumed 0–120% (average 22±26%) of phytoplankton daily growth. Grazing and growth rates found in this study were low compared to rates reported in another Arctic system, the Barents Sea, and in major geographic regions of the world ocean.     

Variability of chlorophyll and primary production in the Eastern North Atlantic Subtropical Gyre: potential factors affecting phytoplankton activity

Size-fractionated chlorophyll-a and carbon incorporation rates were determined on a series of 13 cruises carried out from 1992 to 2001with the aim of investigating the patterns and causes of variability in phytoplankton chlorophyll and production in the Eastern North Atlantic Subtropical Gyral Province (NASE). Averaged (±SE) integrated chlorophyll-a concentration and primary production rate were 17±1 mg m⁻² and 253±22 mg C m⁻² d⁻¹. Small-sized cells (<2 μm) formed the bulk of phytoplankton biomass (71%) and accounted for 54% of total primary production. A clear latitudinal gradient in these variables was not detected. By contrast, large seasonal variability was detected in terms of primary production, although integrated phytoplankton biomass, as estimated from chlorophyll-a concentration, remained rather constant and did not display significant changes with time. Variability in primary production (PP) was related mainly to variability in surface temperature and surface chlorophyll-a concentration. The control exerted by surface temperature was related to nutrient availability. By contrary, euphotic-zone depth, depth of maximum concentration of chlorophyll-a and integrated chlorophyll-a did not contribute significantly to the high variability in primary production observed in this oligotrophic region.     

Basin-scale variability of phytoplankton biomass,production and growth in the Atlantic Ocean

The latitudinal distributions of phytoplankton biomass, composition and production in the Atlantic Ocean were determined along a 10,000-km transect from 50°N to 50°S in October 1995, May 1996 and October 1996. Highest levels of euphotic layer-integrated chlorophyll a (Chl a) concentration (75–125 mg Chl m⁻²) were found in North Atlantic temperate waters and in the upwelling region off NW Africa, whereas typical Chl a concentrations in oligotrophic waters ranged from 20 to 40 mg Chl m⁻². The estimated concentration of surface phytoplankton carbon (C) biomass was 5–15 mg C m⁻² in the oligotrophic regions and increased over 40 mg C m⁻² in richer areas. The deep chlorophyll maximum did not seem to constitute a biomass or productivity maximum, but resulted mainly from an increase in the Chl a to C ratio and represented a relatively small contribution to total integrated productivity. Primary production rates varied from 50 mg C m⁻² d⁻¹ at the central gyres to 500–1000 mg C m⁻² d⁻¹ in upwelling and higher latitude regions, where faster growth rates (μ) of phytoplankton (>0.5 d⁻¹) were also measured. In oligotrophic waters, microalgal growth was consistently slow [surface μ averaged 0.21±0.02 d⁻¹ (mean±SE)], representing <20% of maximum expected growth. These results argue against the view that the subtropical gyres are characterized by high phytoplankton turnover rates. The latitudinal variations in μ were inversely correlated to the changes in the depth of the nitracline and positively correlated to those of the integrated nitrate concentration, supporting the case for the role of nutrients in controlling the large-scale distribution of phytoplankton growth rates. We observed a large degree of temporal variability in the phytoplankton dynamics in the oligotrophic regions: productivity and growth rates varied in excess of 8-fold, whereas microalgal biomass remained relatively constant. The observed spatial and temporal variability in the biomass specific rate of photosynthesis is at least three times larger than currently assumed in most satellite-based models of global productivity.     

Latitudinal distribution of microbial plankton abundance,production, and respiration in the Equatorial Atlantic in autumn 2000

Phytoplankton and bacterial abundance, size-fractionated phytoplankton chlorophyll-a (Chl-a) and production together with bacterial production, microbial oxygen production and respiration rates were measured along a transect that crossed the Equatorial Atlantic Ocean (10°N–10°S) in September 2000, as part of the Atlantic Meridional Transect 11 (AMT 11) cruise. From 2°N to 5°S, the equatorial divergence resulted in a shallowing of the pycnocline and the presence of relatively high nitrate (>1 μM) concentrations in surface waters. In contrast, a typical tropical structure (TTS) was found near the ends of the transect. Photic zone integrated ¹⁴C primary production ranged from ∼200 mg C m⁻² d⁻¹ in the TTS region to ∼1300 mg C m⁻² d⁻¹ in the equatorial divergence area. In spite of the relatively high primary production rates measured in the equatorial upwelling region, only a moderate rise in phytoplankton biomass was observed as compared to nearby nutrient-depleted areas (22 vs. 18 mg Chl-a m⁻², respectively). Picophytoplankton were the main contributors (>60%) to both Chl-a biomass and primary production throughout the region. The equatorial upwelling did not alter the phytoplankton size structure typically found in the tropical open ocean, which suggests a strong top-down control of primary producers by zooplankton. However, the impact of nutrient supply on net microbial community metabolism, integrated over the euphotic layer, was evidenced by an average net microbial community production within the equatorial divergence (1130 mg C m⁻² d⁻¹) three-fold larger than net production measured in the TTS region (370 mg C m⁻² d⁻¹). The entire region under study showed net autotrophic community metabolism, since respiration accounted on average for 51% of gross primary production integrated over the euphotic layer.     

Seasonal and spatial variability in plankton production and respiration in the Subtropical Gyres of the Atlantic Ocean

Euphotic zone plankton production (P) and respiration (R) were determined from the in vitro flux of dissolved oxygen during six latitudinal transects of the Atlantic Ocean, as part of the Atlantic Meridional Transect (AMT) programme. The transects traversed the North and South Atlantic Subtropical Gyres (N gyre, 18–38°N; S gyre, 11–35°S) in April–June and September–November 2003–2005. The route and timing of the cruises enabled the assessment of the seasonal variability of P, R and P/R in the N and S gyres, and the comparison of the previously unsampled N gyre centre with the more frequently sampled eastern edge of the gyre. Mean euphotic zone integrated rates (±SE) were P=63±23 (n=31), R=69±22 (n=30) mmol O₂ m⁻² d⁻¹ in the N gyre; and P=58±26 (n=30), R=62±24 (n=30) mmol O₂ m⁻² d⁻¹ in the S gyre. Overall, the N gyre was heterotrophic (R>P) and it was more heterotrophic than the S gyre, but the metabolic balance of both gyres changed with season. Both gyres were net heterotrophic in autumn, and balanced in spring. This seasonal contrast was most pronounced for the S gyre, because it was more autotrophic than the N gyre during spring. This may have arisen from differences in nitrate availability, because spring sampling in the S gyre coincided with periods of deep mixing to the nitracline, more frequently than spring sampling within the N gyre. Our results indicate that the N gyre is less heterotrophic than previous estimates suggested, and that there is an apparent decrease in R from the eastern edge to the centre of the N gyre, possibly indicative of an allochthonous organic carbon source to the east of the gyre.     

Island mass effect in the Juan Fernández Archipelago (33°S), Southeastern Pacific

Spatial and temporal variability of the island mass effect (IME; defined as local increases of phytoplankton associated with the presence of islands) at the Juan Fernández Archipelago (JFA) is analyzed using chlorophyll-a (Chl-a) satellite data, altimetry, sea surface temperature, wind, geostrophic currents and net heat flux over a ten year period (2002–2012). The the JFA islands (Robinson Crusoe-Santa Clara (RC-SC) and Alejandro Selkirk (AS)) present wakes with significant Chl-a increases, mainly during spring time. These wakes can reach Chl-a values of one order of magnitude higher (~1 mg m⁻³) than the surrounding oligotrophic waters (<0.1 mg m⁻³). The wakes are similar to von Kármán vortex streets which have been used to explain the impact of IME on Chl-a increases in numerical models. The wakes are formed from a high productivity area in the lee of the island, extending to the oceanic region as high Chl-a patches associated with submesoscale eddies that are detached from the islands and connected by less-productive zones. This pattern coincides with previous models that predict the effects of island-generated flow perturbations on biological production variability. The IME is a recurrent feature of islands that has even been observed in decadal average fields. In such average fields, the Chl-a values in RC-SC and AS islands can exceed values found in a Control Zone (a zone without islands) by ~50% and 30%, respectively. Seasonal and interannual variability reveals that, as a consequence of the IME, the winter Chl-a maximum associated with the development of winter convection and mesoscale eddies that propagate from the continental zone, promote that the Chl-a maximum extends towards spring. The IME has an impact on the island on both a local as well as a more regional scale that affects an area of ~40,000 km² (1°Latitude×4°Longitude) centered on the islands. The transport of high productivity patches associated with submesoscale eddies may be responsible for IME propagation at a regional scale. Around the islands, the presence of a weak oceanic incident flow and strong and recurrent wind-wakes, suggest that the generation of Chl-a wakes result from a combined effect between both forcings.     

Carbon fluxes through major phytoplankton groups during the spring bloom and post-bloom in the Northwestern Mediterranean Sea

The carbon flux through major phytoplankton groups, defined by their pigment markers, was estimated in two contrasting conditions of the Northwestern Mediterranean open ocean ecosystem: the spring bloom and post-bloom situations (hereafter Bloom and Post-bloom, respectively). During Bloom, surface chlorophyll a (Chl a) concentration was higher and dominated by diatoms (53% of Chl a), while during Post-bloom Synechococcus (42%) and Prymnesiophyceae (29%) became dominant. The seawater dilution technique, coupled to high pressure liquid chromatography (HPLC) analysis of pigments and flow cytometry (FCM), was used to estimate growth and grazing rates of major phytoplankton groups in surface waters. Estimated growth rates were corrected for photoacclimation based on FCM-detected changes in red fluorescence per cell. Given the 30% average decrease in the pigment content per cell between the beginning and the end of the incubations, overlooking photoacclimation would have resulted in a 0.40 d^?1 underestimation of phytoplankton growth rates. Corrected average growth rates (μ_o) were 0.90±0.20 (SD) and 0.40±0.14 d^?1 for Bloom and Post-bloom phytoplankton, respectively. Diatoms, Cryptophyceae and Synechococcus were identified as fast-growing groups and Prymnesiophyceae and Prasinophyceae as slow-growing groups across Bloom and Post-bloom conditions. The higher growth rate during Bloom was due to dominance of phytoplankton groups with higher growth rates than those dominating in Post-bloom. Average grazing rates (m) were 0.58±0.20 d^?1 (SD) and 0.31±0.07 d^?1. The proportion of phytoplankton growth consumed by microzooplankton grazing (m/μ_o) tended to be lower in Bloom (0.69±0.34) than in Post-bloom (0.80±0.08). The intensity of nutrient limitation experienced by phytoplankton indicated by μ_o/μ_n (where μ_n is the nutrient-amended growth rate), was similar during Bloom (0.78) and Post-bloom (0.73). Primary production from surface water (PP) was estimated with ¹⁴C incubations. A combination of PP and Chl a synthesis rate yielded C/Chl a ratios of 34±21 and 168±75 (g:g) for Bloom and Post-bloom, respectively. Transformation of group-specific Chl a fluxes into carbon equivalents confirmed the dominant role of diatoms during Bloom and Synechococcus and Prymnesiophyceae during Post-bloom.     

Top-down control of spring surface phytoplankton blooms by microzooplankton in the Central Yellow Sea,China

Phytoplankton growth and microzooplankton grazing were studied during the 2007 spring bloom in Central Yellow Sea. The surveyed stations were divided to pre-bloom phase (Chl a concentration less than 2 μg L⁻¹), and bloom phase (Chl a concentration greater than 2 μg L⁻¹). Shipboard dilution incubation experiments were carried out at 19 stations to determine the phytoplankton specific growth rates and the specific grazing rates of microzooplankton on phytoplankton. Diatoms dominated in the phytoplankton community in surface waters at most stations. For microzooplankton, Myrionecta rubra and tintinnids were dominant, and heterotrophic dinoflagellate was also important in the community. Phytoplankton-specific growth rates, with an average of 0.60±0.19 d⁻¹, were higher at pre-bloom stations (average 0.62±0.17 d⁻¹), and lower at the bloom stations (average 0.59±0.21 d⁻¹), but the difference of growth rates between bloom and pre-bloom stations was not statistically significant (t test, p=0.77). The phytoplankton mortality rate by microzooplankton grazing averaged 0.41±0.23 d⁻¹ at pre-bloom stations, and 0.58±0.31 d⁻¹ during the blooms. In contrast to the growth rates, the statistic difference of grazing rates between bloom and pre-bloom stations was significant (after removal of outliers, t test, p=0.04), indicating the importance of the top-down control in the phytoplankton bloom processes. Average potential grazing efficiency on primary productivity was 66% at pre-bloom stations and 98% at bloom stations, respectively. Based on our results, the biomass maximum phase (bloom phase) was not the maximum growth rate phase. Both phytoplankton specific growth rate and net growth rate were higher in the pre-bloom phase than during the bloom phase. Microzooplankton grazing mortality rate was positively correlated with phytoplankton growth rate during both phases, but growth and grazing were highly coupled during the booming phase. There was no correlation between phytoplankton growth rate and cell size during the blooms, but they were positive correlated during the pre-bloom phase. Our results indicate that microzooplankton grazing is an important process controlling the growth of phytoplankton in spring bloom period in the Central Yellow Sea, particularly in the “blooming” phase.     

Phytoplankton growth and mortality during the 1995 Northeast Monsoon and Spring Intermonsoon in the Arabian Sea

Phytoplankton growth rates and mortality rates were experimentally examined at eight stations in the Arabian Sea along the U.S. JGOFS cruise track during the 1995 Northeast Monsoon (January) and Spring Intermonsoon (March–April). Instantaneous growth rates averaged over an entire cruise were approximately twice as high during the NE Monsoon than during the Spring Intermonsoon period (overall averages of 0.84±0.29 (s.d.) versus 0.44±0.19 d⁻¹). Average herbivore grazing (mortality) rates, however, were quite similar for the two seasons (overall averages of 0.35±0.18 and 0.30±0.17 d⁻¹ for the NE Monsoon and Spring Intermonsoon, respectively). The absolute amounts of phytoplankton biomass consumed during each season also were similar (29 and 25% of standing stock consumed d⁻¹ for the January and March–April cruises, respectively), as were the geographical trends of this removal. These seasonal trends in growth and removal rates resulted in net phytoplankton growth rates that were considerably higher during the January cruise (0.48 d⁻¹) than during the March–April cruise (0.14 d⁻¹). That is, phytoplankton production was more closely balanced during the Spring Intermonsoon season (87% of daily primary production consumed) relative to the NE Monsoon season (49% of daily primary production consumed). Station-to-station variability was high for rate measurements during either cruise. Nevertheless, there was a clear onshore–offshore trend in the absolute rate of removal of phytoplankton biomass (μg chlorophyll consumed l⁻¹ d⁻¹) during both cruises. Coastal stations had removal rates that were typically 2–4 times higher than removal rates at oceanic stations.     

Diatom biomass and productivity in oceanic and plume-influenced waters of the western tropical Atlantic ocean

Whereas diatoms (class Bacillariophyceae) often dominate phytoplankton taxa in the Amazon estuary and shelf, their contribution to phytoplankton dynamics and impacts on regional biogeochemistry are poorly understood further offshore in the western tropical Atlantic Ocean (WTAO). Thus, relative contribution of diatoms to phytoplankton biomass and primary production rates and associated environmental conditions were quantified during three month-long cruises in January–February 2001, July–August 2001, and April–May 2003. The upper water column was sampled at 6 light depths (100%, 50%, 25%, 10%, 1% and 0.1% of surface irradiance) at 64 stations between 3° and 14°N latitude and 41° and 58°W longitude. Each station was categorized as ‘oceanic’ or ‘plumewater’, based on principal component analysis of eight physical, chemical and biological variables. All stations were within the North Brazil Current, and plumewater stations were characterized by shallower mixed layers with lower surface salinities and higher dissolved silicon (dSi) concentrations than oceanic stations. The major finding was a much greater role of diatoms in phytoplankton biomass and productivity at plumewater stations relative to oceanic stations. Mean depth-integrated bSi concentrations at the plumewater and oceanic stations were 14.2 and 3.7 mmol m⁻², respectively. Mean depth-integrated SiP rates at the plumewater and oceanic stations were 0.17 and 0.02 mmol m⁻² h⁻¹, respectively. Based on ratios of SiP and PP rates, and typical Si:C ratios, diatoms contributed on average 29% of primary productivity at plumewater stations and only 3% of primary productivity at oceanic stations. In contrast, phytoplankton biomass (as chlorophyll a concentrations) and primary production (PP) rates (as ¹⁴C uptake rates) integrated over the euphotic zone were not significantly different at plumewater and oceanic stations. Chlorophyll a concentrations ranged from 8.5 to 42.4 mg m⁻² and 4.0 to 38.0 mg m⁻² and PP rates ranged from 2.2 to 11.2 mmol m⁻² h⁻² and 1.8 to 10.8 mmol m⁻² h⁻² at plumewater and oceanic stations, respectively. A conservative estimate of annual integrated SiP in offshore waters of Amazon plume between April and August is 0.59 Tmol Si, based on mean SiP rates in plumewaters and satellite-derived estimates of the area of the Amazon plume. In conclusion, river plumewaters dramatically alter the silicon dynamics of the WTAO, forming extensive diatom-dominated phytoplankton blooms that may contribute significantly to the global Si budget as well as contributing to energy and matter flow off of the continental shelf.     

Light absorption by phytoplankton,non-algal particles and dissolved organic matter at the Patagonia shelf-break in spring and summer

Satellite image studies and recent in situ sampling have identified conspicuous phytoplankton blooms during spring and summer along the Patagonia shelf-break front. The magnitudes and spectral characteristics of light absorption by total particulate matter (phytoplankton and detritus) and colored dissolved organic matter (CDOM) have been determined by spectrophotometry in that region for spring 2006 and late summer 2007 seasons. In spring, phytoplankton absorption was the dominant optical component of light absorption (60–85%), and CDOM showed variable and important contributions in summer (10–90%). However, there was a lack of correlation between phytoplankton biomass (chlorophyll-a concentration or [chl a]) and the non-algal compartment in both periods. A statistically significant difference was found between the two periods with respect to the CDOM spectral shape parameter (S_cdom), with means of 0.015 (spring) and 0.012 nm^?1 (summer). Nonetheless, the mean S_cdm values, which describe the slope of detritus plus CDOM spectra, did not differ between the periods (average of 0.013 nm^?1). Phytoplankton absorption values in this work showed deviations from mean parameterizations in previous studies, with respect to [chl a], as well as between the two study periods. In spring, despite the microplankton dominance, high specific absorption values and large dispersion were found (a*_ph(440)=0.04±0.03 m² mg [chl a]^?1), which could be attributed to an important influence of photo-protector accessory pigments. In summer, deviations from general trends, with values of a*_ph(440) even higher (0.09±0.02 m² mg [chl a]^?1), were due to the dominance of small cell sizes and also to accessory pigments. These results highlight the difficulty in deriving robust relationships between chlorophyll concentration and phytoplankton absorption coefficients regardless of the season period. The validity of a size parameter (S_f) derived from the absorption spectra has been demonstrated and was shown to describe the size structure of phytoplankton populations, independently of pigment concentration, with mean values of 0.41 in spring and 0.72 in summer. Our results emphasize the need for specific parameterization for the study region and seasonal sampling approach in order to model the inherent optical properties from water reflectance signatures.     

Factors influencing particulate lipid production in the East Atlantic Ocean

Extensive analyses of particulate lipids and lipid classes were conducted to gain insight into lipid production and related factors along the biogeochemical provinces of the Eastern Atlantic Ocean. Data are supported by particulate organic carbon (POC), chlorophyll a (Chl a), phaeopigments, Chl a concentrations and carbon content of eukaryotic micro-, nano- and picophytoplankton, including cell abundances for the latter two and for cyanobacteria and prokaryotic heterotrophs. We focused on the productive ocean surface (2 m depth and deep Chl a maximum (DCM)). Samples from the deep ocean provided information about the relative reactivity and preservation potential of particular lipid classes. Surface and DCM particulate lipid concentrations (3.5–29.4 μg L⁻¹) were higher than in samples from deep waters (3.2–9.3 μg L⁻¹) where an increased contribution to the POC pool was observed. The highest lipid concentrations were measured in high latitude temperate waters and in the North Atlantic Tropical Gyral Province (13–25°N). Factors responsible for the enhanced lipid synthesis in the eastern Atlantic appeared to be phytoplankton size (micro, nano, pico) and the low nutrient status with microphytoplankton having the most expressed influence in the surface and eukaryotic nano- and picophytoplankton in the DCM layer. Higher lipid to Chl a ratios suggest enhanced lipid biosynthesis in the nutrient poorer regions. The various lipid classes pointed to possible mechanisms of phytoplankton adaptation to the nutritional conditions. Thus, it is likely that adaptation comprises the replacement of membrane phospholipids by non-phosphorus containing glycolipids under low phosphorus conditions. The qualitative and quantitative lipid compositions revealed that phospholipids were the most degradable lipids, and their occurrence decreased with increasing depth. In contrast, wax esters, possibly originating from zooplankton, survived downward transport probably due to the fast sinking rate of particles (fecal pellets). The important contribution of glycolipids in deep waters reflected their relatively stable nature and degradation resistance. A lipid-based proxy for the lipid degradative state (Lipolysis Index) suggests that many lipid classes were quite resistant to degradation even in the deep ocean.     

Distribution of bacterial abundance and cell-specific nucleic acid content in the Northeast Pacific Ocean

We tested the idea that bacterial cells with high nucleic acid content (HNA cells) are the active component of marine bacterioplankton assemblages, while bacteria with low nucleic acid content (LNA cells) are inactive, with a large data set (>1700 discrete samples) based on flow cytometric analysis of bacterioplankton in the Northeast Pacific Ocean off the coast of Oregon and northern California, USA. Samples were collected in the upper 150 m of the water column from the coast to 250 km offshore during 14 cruises from March 2001 to September 2003. During this period, a wide range of trophic states was encountered, from dense diatom blooms (chlorophyll-a concentrations up to 43 μg l⁻¹) at shelf stations during upwelling season (March–September) to lower chlorophyll-a concentrations (0.1–5 μg l⁻¹) during winter (November–February) and at basin stations (>1700 m depth). We found only weakly positive relations of log total bacterial abundance to log chlorophyll-a concentration (as a proxy for availability of organic substrate), and of HNA bacteria as a fraction of total bacteria to log chlorophyll-a. Abundance of HNA and LNA bacteria co-varied positively in all regions, although HNA bacteria were more responsive to high phytoplankton biomass in shelf waters than in slope and basin waters. Since LNA cell abundance in general showed responses similar to those of HNA cell abundance to changes in phytoplankton biomass, our data do not support the hypothesis that HNA cells are the sole active component of marine bacterioplankton.     

Modelling phytoplankton deposition to Chesapeake Bay sediments during winter–spring: interannual variability in relation to river flow

The often-rapid deposition of phytoplankton to sediments at the end of the spring phytoplankton bloom is an important component of benthic–pelagic coupling in temperate and high latitude estuaries and other aquatic systems. However, quantifying the flux is difficult, particularly in spatially heterogeneous environments. Surficial sediment chlorophyll-a, which can be measured quickly at many locations, has been used effectively by previous studies as an indicator of phytoplankton deposition to estuarine sediments. In this study, surficial sediment chlorophyll-a was quantified in late spring at 20–50 locations throughout Chesapeake Bay for 8 years (1993–2000). A model was developed to estimate chlorophyll-a deposition to sediments using these measurements, while accounting for chlorophyll-a degradation during the time between deposition and sampling. Carbon flux was derived from these estimates via C:chl-a = 75.Bay-wide, the accumulation of chlorophyll-a on sediments by late spring averaged 171 mg m⁻², from which the chlorophyll-a and carbon sinking fluxes, respectively, were estimated to be 353 mg m⁻² and 26.5 gC m⁻². These deposition estimates were ∼50% of estimates based on a sediment trap study in the mid-Bay. During 1993–2000, the highest average chlorophyll-a flux was in the mid-Bay (248 mg m⁻²), while the lowest was in the lower Bay (191 mg m⁻²). Winter–spring average river flow was positively correlated with phytoplankton biomass in the lower Bay water column, while phytoplankton biomass in that same region of the Bay was correlated with increased chlorophyll-a deposition to sediments. Responses in other regions of the Bay were less clear and suggested that the concept that nutrient enrichment in high flow years leads to greater phytoplankton deposition to sediments may be an oversimplification. A comparison of the carbon flux associated with the deposition of the spring bloom with annual benthic carbon budgets indicated that the spring bloom did not contribute a disproportionately large fraction of annual carbon inputs to Chesapeake Bay sediments. Regional patterns in chlorophyll-a deposition did not correspond with the strong regional patterns that have been found for plankton net community metabolism during spring.     

Nutrient and phytoplankton dynamics off the west coast of Vancouver Island during the 1997/98 ENSO event

Six research cruises were conducted off the west coast of Vancouver Island between April and October of 1997 and 1998 as part of the Canadian GLOBEC project to compare nutrient and phytoplankton dynamics between ENSO (1997) and non-ENSO (1998) years. Limited sampling also was conducted during three cruises in 1999. During the 1997 ENSO period, there was a shallow thermocline (∼10 m) that resulted in a shallower mixed layer, lower salinity and density, and stronger summer stratification. In general on the shelf, the 1997 growing season was characterized by higher nitrate (7.5 μM) and silicic acid (17 μM) concentrations, lower total chlorophyll (∼76 mg m⁻²), lower phytoplankton carbon biomass (0.2 mg C L⁻¹), and lower diatom abundance and biomass than in 1998. Phytoplankton assemblages were dominated by nanoplankton in 1997 and by diatoms in 1998. These results suggest that the 1997 ENSO was responsible for a reduction in the growth and biomass of larger phytoplankton cells. In mid-1998, the hydrographic characteristics off the west coast of Vancouver Island changed suddenly. The 1997 poleward transport of warm water reversed to an equatorward transport of coastal water in July 1998, which was accompanied by normal summer upwelling. During 1998, a large diatom bloom (mainly dominated by Chaetoceros debilis, Leptocylindrus danicus and to a lesser extent by Skeletomema and Pseudo-nitzschia sp.) was observed in July over the continental shelf. This large bloom resulted in chlorophyll concentrations of up to 400 mg m⁻², primary productivity of up to 11 g C m⁻² d⁻¹, and near undetectable dissolved nitrogen concentrations at some of the shelf stations in 1998. In contrast, during 1997, the sub-tropical waters that were advected over the slope, resulted in low chlorophyll a and primary productivity (generally <1 g C m⁻² d⁻¹). Therefore, there was a sharp contrast between the very high primary productivity on the shelf in July 1998, due to normal nutrient replenishment from summer upwelling and outflow from the Strait of Juan de Fuca, and the lower primary productivity during the 1997 ENSO year. During 1998, non-ENSO conditions resulted in phytoplankton biomass that was twice as high on the shelf as that measured in regions beyond the continental shelf of the west coast of Vancouver Island.     

Microbial biomass and viral infections of heterotrophic prokaryotes in the sub-surface layer of the central Arctic Ocean

Seawater samples were collected for microbial analyses between 55 and 235 m depth across the Arctic Ocean during the SCICEX 97 expedition (03 September–02 October 1997) using a nuclear submarine as a research platform. Abundances of prokaryotes (range 0.043–0.47×10⁹ dm⁻³) and viruses (range 0.68–11×10⁹ dm⁻³) were correlated (r=0.66, n=150) with an average virus:prokaryote ratio of 26 (range 5–70). Biomass of prokaryotes integrated from 55 to 235 m ranged from 0.27 to 0.85 g C m⁻² exceeding that of phytoplankton (0.005–0.2 g C m⁻²) or viruses (0.02–0.05 g C m⁻²) over the same depth range by an order of magnitude on average. Using transmission electron microscopy (TEM), we estimated that 0.5% of the prokaryote community on average (range 0–1.4%) was visibly infected with viruses, which suggests that very little of prokaryotic secondary production was lost due to viral lysis. Intracellular viruses ranged from 5 to >200/cell, with an average apparent burst size of 45±38 (mean±s.d.; n=45). TEM also revealed the presence of putative metal-precipitating bacteria in 8 of 13 samples, which averaged 0.3% of the total prokaryote community (range 0–1%). If these prokaryotes are accessible to protistan grazers, the Fe and Mn associated with their capsules might be an important source of trace metals to the planktonic food web. After combining our abundance and mortality data with data from the literature, we conclude that the biomass of prokaryoplankton exceeds that of phytoplankton when averaged over the upper 250 m of the central Arctic Ocean and that the fate of this biomass is poorly understood.     

Relationships between picophytoplankton and optical properties in the Azores Front region in the Atlantic Ocean

The picophytoplankton evolved to become extremely effective harvesters of light and, thus, dominating productivity in the open ocean. In this study, their distribution in relation to the underwater light field was examined in the Azores Front region of the North Atlantic. In this region, attenuation coefficients of downward irradiance varied between 0.038 and 0.065 m^− 1. Maximum absorptions were at the red and green parts of the light spectrum, typical of the oligotrophic ocean. The euphotic zone ranged from 70 to 120 m. The deep chlorophyll maximum (DCM) was found at depths where subsurface light ranged between 0.1 and 1% of its surface values. Prochlorococcus was the dominant phytoplankton group while Synechococcus (cyanobacteria) and picoeukaryotes were much less abundant. The ability of chlorophyll to absorb light, i.e., the absorption coefficient of chlorophyll (a^⁎), was found to be dependent on the photoacclimation of the cells and was lower at low-light intensities. Due to a packaging effect and probable pigment changes, a^⁎ at the DCM was the lowest while the chlorophyll per Prochlorococcus cell was the highest. This is a major adaptation of the picophytoplankton to low light (less than 1% surface light), which allows them to bloom at the DCM. This study indicates that the Atlantic Ocean models for the estimation of phytoplankton concentration and/or primary production should not use a constant carbon biomass-to-chlorophyll ratio for phytoplankton based on phytoplankton size, and should take into account the variation in chlorophyll-absorption ability.