Vertical distributions of large ontogenetically migrating copepods in the Oyashio region during their growing season

Neocalanus flemingeri, Neocalanus plumchrus, Neocalanus cristatus and Eucalanus bungii are large and dominant mesozooplankton occurring throughout the subarctic Pacific. They are an important trophic link and transporter of organic matter to the mesopelagic zone. Vertical distributions of these copepods were investigated from March to October 2000 in the Oyashio region of the western subarctic Pacific. Neocalanus plumchrus and N. flemingeri were distributed in the surface layer (0–50 m) and N. cristatus and E. bungii in the subsurface layer (50–100 m). However, when examined in detail, clear seasonal and vertical differences were observed. Neocalanus plumchrus was concentrated in the top 20 m from late April to the end of July, and N. flemingeri showed a little deeper distribution from May to July. Neocalanus cristatus showed a deeper distribution than that of grazing individuals of E. bungii from April to early July, but grazing individuals of E. bungii (C3–C6) showed a deeper distribution than that of N. cristatus from the end of July to October. Early copepodites of E. bungii were distributed much shallower than late copepodite stages and overlapped with copepodites of N. plumchrus and N. flemingeri. These results suggest that the four species of large copepods have established habitat segregation by season, vertical distribution and food resource partitioning in the Oyashio region as well as other regions of the North Pacific.     

Meso- and microzooplankton responses to an in situ iron fertilization experiment (SEEDS II) in the northwest subarctic Pacific

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

Mesozooplankton responses to iron-fertilization in the western subarctic Pacific (SEEDS2001)

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

Seasonal changes in zooplankton abundance,biomass, size structure and dominant copepods in the Oyashio region analysed by an optical plankton counter

To identify seasonal patterns of change in zooplankton communities, an optical plankton counter (OPC) and microscopic analysis were utilised to characterise zooplankton samples collected from 0 to 150 m and 0 to 500 m in the Oyashio region every one to three months from 2002 to 2007. Based on the OPC measurements, the abundance and biomass of zooplankton peaked in June (0–150 m) or August (150–500 m), depending on the depth stratum. The peak periods of the copepod species that were dominant in terms of abundance and biomass indicated species-specific patterns. Three Neocalanus species (Neocalanus cristatus, Neocalanus flemingeri and Neocalanus plumchrus) exhibited abundance peaks that occurred before their biomass peaks, whereas Eucalanus bungii and Metridia pacifica experienced biomass peaks before their abundance peaks. The abundance peaks corresponded to the recruitment periods of early copepodid stages, whereas the biomass peaks corresponded to the periods when the dominant populations reached the late copepodid stages (C5 or C6). Because the reproduction of Neocalanus spp. occurred in the deep layer (>500 m), their biomass peaks were observed when the major populations reached stage C5 after the abundance peaks of the early copepodid stages. The reproduction of E. bungii and M. pacifica occurred near the surface layer. These species first formed biomass peaks of C6 and later developed abundance peaks of newly recruited early copepodid stages. From the comparison between OPC measurements and microscopic analyses, seasonal changes in zooplankton biomass at depths of 0–150 m were governed primarily by E. bungii and M. pacifica, whereas those at depths of 150–500 m were primarily caused by the three Neocalanus species.     

Particulate suspended matter concentrations in the Bahía Blanca Estuary,Argentina: Implication for the development of phytoplankton blooms

The inner zone of the Bahía Blanca Estuary is shallow, nutrient-rich and turbid. Tidal energy and water turbulence strongly affect the water column resulting in a well-mixed structure and high concentrations of suspended sediment. The phytoplankton community is mostly dominated by diatoms and the annual pattern has been characterized by a recurrent winter-early spring bloom. Here, we investigated to what extent the temporal variations of suspended particulate matter (SPM) regulate the phytoplankton blooms in the head of the estuary by light-limitation. Sampling was done on a fortnightly basis (weekly during the blooming season) at a fixed station in the inner zone of the estuary from January 2007 to February 2008. SPM concentrations and light extinction coefficients (k) in the water column were significantly correlated and showed relatively lower values during the phytoplankton maximal biomass levels. During winter, SPM and k reached values of 23.6 mg l⁻¹ and 0.17 m⁻¹ which were significantly lower than the annual means of 77.6 mg l⁻¹ and 2.94 m⁻¹, respectively. The particulate organic matter (POM) concentration was significantly correlated with the calculated phytoplankton biomass although the contribution of the latter to the total POM was rather low. Both, POM and biomass, had maximal values during winter (21.8 mg l⁻¹ and 393.5 μg C l⁻¹) and mid summer (24.3 mg l⁻¹ and 407.0 μg C l⁻¹), with cell densities up to 8 × 10⁶ cells l⁻¹ and chlorophyll a up to 24.6 μg l⁻¹. Our results suggest that the decrease of SPM concentrations in the water column with a concomitant increase in the penetration of solar radiation seems to be one of the main causes for the development of the phytoplankton winter bloom in the Bahía Blanca Estuary.     

Lagrangian ensemble model of Copepoda (Neocalanus cristatus) in the northwestern subarctic Pacific

A Lagrangian ensemble model describing the population dynamics of Neocalanus cristatus was developed. To describe the ecology of N. cristatus, life stage must be included in the model. For this purpose, a new zooplankton compartment, divided into nine life stages, was added to the North Pacific ecosystem model for understanding regional oceanography. In addition, we introduced Lagrangian particles for the copepod population. Each Lagrangian particle represented a population of the same cohort of copepods with information on representative age, developmental stage, growth rate, structural weight, and food satiation. The new model revealed that surviving cohorts are only matched with the phytoplankton bloom after the nauplii stages. The model showed that the existing phytoplankton concentration when copepodites of N. cristatus appear in the surface water determines the development pattern of copepodite stages. The timing of the seasonal phytoplankton bloom depends on climate change, and a sensitivity analysis showed that the multiple spawning strategy through time has an important role in matching the unstable seasonal bloom cycle on a daily basis. The model including the cohorts can collaborate closely with weekly or monthly observations to reveal the rapid response of the marine ecosystem.     

Iron deficiency in micro-sized diatoms in the Oyashio region of the Western subarctic Pacific during spring

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

Plankton community diversity from bacteria to copepods in bloom and non-bloom conditions in the Celtic Sea in spring

The plankton community composition comprising heterotrophic bacteria, pro-/eukaryotes, heterotrophic nanoflagellates, microzooplankton and mesozooplankton was assessed during the spring bloom and at non-bloom stations in the English Channel and Celtic Sea between 6 and 12 April 2002. Non-bloom sites were characterised by a dominance of pro-/eukaryotic phytoplankton <20 μm, higher abundance of heterotrophic nanoflagellates, microzooplankton standing stocks ranging between 60 and 380 mg C m⁻², lower mesozooplankton diversity and copepod abundance of between 760 and 2600 ind m⁻³. Within the bloom, the phytoplankton community was typically dominated by larger cells with low abundance of pro-/eukaryotes. Heterotrophic nanoflagellate cell bio-volume decreased leading to a reduction in biomass whereas microzooplankton biomass increased (360–1500 mg C m⁻²) due to an increase in cell bio-volume and copepod abundance ranged between 1400 and 3800 ind m⁻³. Mesozooplankton diversity increased with an increase in productivity. Relationships between the plankton community and environmental data were examined using multivariate statistics and these highlighted significant differences in the abiotic variables, the pro-/eukaryotic phytoplankton communities, heterotrophic nanoflagellate, microzooplankton and total zooplankton communities between the bloom and non-bloom sites. The variables which best described variation in the microzooplankton community were temperature and silicate. The spatial variation in zooplankton diversity was best explained by temperature. This study provides an insight into the changes that occur between trophic levels within the plankton in response to the spring bloom in this area.     

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.     

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.     

Community Structure and Dynamics of Phytoplankton in the Western Subarctic Pacific Ocean: A Synthesis

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

Functional roles of interzonal migrating mesozooplankton in the western subarctic Pacific

Grazing experiments and production estimation based on life-history analysis of Neocalanus copepods (N. cristatus, N. plumchrus and N. flemingeri) were carried out in the Oyashio region to understand the carbon flows associated with the interzonal migrating copepods. These copepods, and also Eucalanus bungii, fed on nano- and micro-sized organisms non-selectively throughout the season. However, diatoms were the dominant food resource until May and organisms, such as ciliates were the major resource after May. Daily growth rate was estimated from the Ikeda–Motoda, Huntley–Lopez and Hirst–Sheader models. Since the growth rates were considered to be overestimates for the Huntley–Lopez model and underestimates for the other two models, we applied the weight-specific growth rates previously reported for these species in the Bering Shelf. Surface biomass of Neocalanus increased rapidly in June during the appearance of C5, and a successive increase of overwintering stock was evident in the deeper layer. The deep biomass decreased gradually from September to May during the dormant and reproduction period. N. cristatus has the largest annual mean biomass (2.3 gC m⁻²), followed by N. plumchrus (1.1) and N. flemingeri (0.4). Daily production rate of Neocalanus varied from 0.4 to 363.4 mgC m⁻² day⁻¹, to which N. cristatus was the largest contributor. Annual production was estimated as 11.5 gC m⁻² year⁻¹ for N. cristatus, 5.7 for N. plumchrus and 2.1 for N. flemingeri, yielding annual P/B ratio of 5 for each species. The annual production of Neocalanus accounted for 13.2% of the primary production in the Oyashio region. Their fecal pellets were estimated to account for 14.9% (0.7 gC m⁻² year⁻¹) of sinking flux of organic carbon at 1000-m depth. Moreover, their export flux by ontogenetic vertical migration, which is not measured by sediment trap observations, is estimated to be 91.5% (4.3 gC m⁻² year⁻¹) of carbon flux of sinking particles at 1000-m depth. These results suggest the important role of interzonal migrating copepods in the export flux of carbon.     

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.     

The fate of production in the central Arctic Ocean - top-down regulation by zooplankton expatriates?

We estimated primary and bacterial production, mineral nutrients, suspended chlorophyll a (Chl), particulate organic carbon (POC) and nitrogen (PON), abundance of planktonic organisms, mesozooplankton fecal pellet production, and the vertical flux of organic particles of the central Arctic Ocean (Amundsen basin, 89-88° N) during a 3 week quasi-Lagrangian ice drift experiment at the peak of the productive season (August 2001). A visual estimate of ≈15% ice-free surface, plus numerous melt ponds on ice sheets, supported a planktonic particulate primary production of 50-150 mg C m⁻² d⁻¹ (mean 93 mg C m⁻² d⁻¹, n = 7), mostly confined to the upper 10 m of the nutrient replete water column. The surface mixed layer was separated from the rest of the water column by a strong halocline at 20 m depth. Phototrophic biomass was low, generally 0.03-0.3 mg Chl m⁻³ in the upper 20 m and <0.02 mg Chl m⁻³ below, dominated by various flagellates, dinoflagellates and diatoms. Bacterial abundance (typically 3.7-5.3 × 10⁵, mean 4.1 × 10⁵ cells ml⁻¹ in the upper 20 m and 1.3-3.7 × 10⁵, mean 1.9 × 10⁵ cells ml⁻¹ below) and Chl concentrations were closely correlated (r = 0.75). Mineral nutrients (3 μmol NO₃ l⁻¹, 0.45 μmol PO₄ l⁻¹, 4-5 μmol SiO₄ l⁻¹) were probably not limiting the primary production in the upper layer. Suspended POC concentration was ∼30-105 (mean 53) mg C m⁻³ and PON ∼5.4-14.9 (mean 8.2) mg N m⁻³ with no clear vertical trend. The vertical flux of POC in the upper 30-100 m water column was ∼37-92 (mean 55) mg C m⁻² d⁻¹ without clear decrease with depth, and was quite similar at the six investigated stations. The mesozooplankton biomass (≈2 g DW m⁻², mostly in the upper 50 m water column) was dominated by adult females of the large calanoid copepods Calanus hyperboreus and Calanus glacialis (≈1.6 g DW m⁻²). The grazing of these copepods (estimated via fecal pellet production rates) was ≈15 mg C m⁻² d⁻¹, being on the order of 3% and 20% of the expected food-saturated ingestion rates of C. hyperboreus and C. glacialis, respectively. The stage structure of these copepods, dominated by adult females, and their unsatisfied grazing capacity during peak productive period suggest allochthonous origin of these species from productive shelf areas, supported by their long life span and the prevailing surface currents in the Arctic Ocean. We propose that the grazing capacity of the expatriated mesozooplankton population would match the potential seasonal increase of primary production in the future decreased ice perspective, diminishing the likelihood of algal blooms.     

Growth and grazing rate dynamics of major phytoplankton groups in an oligotrophic coastal site

There has been more attention to phytoplankton dynamics in nutrient-rich waters than in oligotrophic ones thus requiring the need to study the dynamics and responses in oligotrophic waters. Accordingly, phytoplankton community in Blanes Bay was overall dominated by Prymnesiophyceae, remarkably constant throughout the year (31 ± 13% Total chlorophyll a, Tchl a) and Bacillariophyta with a more episodic appearance (20 ± 23% Tchl a). Prasinophyceae and Synechococcus contribution became substantial in winter (Prasinophyceae = 30% Tchl a) and summer (Synechococcus = 35% Tchl a). Phytoplankton growth and grazing mortality rates for major groups were estimated by dilution experiments in combination with high pressure liquid chromatography and flow cytometry carried out monthly over two years. Growth rates of total phytoplankton (range = 0.30–1.91 d⁻¹) were significantly higher in spring and summer (μ > 1.3 d⁻¹) than in autumn and winter (μ ∼ 0.65 d⁻¹) and showed a weak dependence on temperature but a significant positive correlation with day length. Microzooplankton grazing (range = 0.03–1.4 d⁻¹) was closely coupled to phytoplankton growth. Grazing represented the main process for loss of phytoplankton, removing 60 ± 34% (±SD) of daily primary production and 70 ± 48% of Tchl a stock. Chla synthesis was highest during the Bacillarophyceae-dominated spring bloom (Chl a_synt = 2.3 ± 1.6 μg Chl a L⁻¹ d⁻¹) and lowest during the following post-bloom conditions dominated by Prymnesiophyceae (Chl a_synt = 0.23 ± 0.08 μg Chl a L⁻¹ d⁻¹). This variability was smoothed when expressed in carbon equivalents mainly due to the opposite dynamics of C:chl a (range = 11–135) and chl a concentration (range = 0.07–2.0 μg chl a L⁻¹). Bacillariophyta and Synechococcus contribution to C fluxes was higher than to biomass because of their fast-growth rate. The opposite was true for Prymnesiophyceae.     

Dynamics and sources of suspended particulate organic matter in the Marennes-Oléron oyster farming bay: Insights from stable isotopes and microalgae ecology

The aim of this study was to distinguish between sources of the complex variety of Marennes-Oléron Bay suspended particulate organic matter (SPOM) contributing to the tropho-dynamics of the Marennes-Oléron oyster farming bay. Basic biomarkers (Chl a, C/N and POC/Chl a ratios), carbon and nitrogen stable isotopes from SPOM were analyzed and the microalgae community was characterized. The sampling strategy was bimonthly from March 2002 to December 2003; samples were taken from an intertidal mudflat. Four main sources contributed to the SPOM pool: terrigenous input from rivers, neritic phytoplankton, resuspended microphytobenthos and periodic inputs from intertidal Zostera noltii meadows. Seasonal fluctuations were observed in both years of the study period: (1) SPOM collected in the spring of 2002 (δ¹³C = −25‰ to −23‰) was mainly composed of fresh estuarine inputs; (2) SPOM from the summer and fall of 2002 and 2003 was predominantly neritic phytoplankton (δ¹³C = − 22‰ to −19‰); (3) SPOM from the winter of 2002, spring of 2003 and winter of 2003 (δ¹³C = −21 to −23‰) was composed of a mixture of decayed terrigenous river inputs and pelagic phytoplankton, which was predominantly resuspended microphytobenthos. In the summer of 2003—the warmest summer on record in southern France and Europe—SPOM was particularly enriched for ¹³C, with δ¹³C values ranging from −14‰ to −12‰. Pulses in δ¹³C values, indicative of ¹³C-enriched decaying materials, extended into the fall. These were attributed to benthic intertidal inputs, including both resuspended microphytobenthos and Z. noltii detritus. Changes in SPOM sources in Marennes-Oléron Bay may lead to differences in the quality of the trophic environment available for reared oysters.     

Close coupling between ammonium uptake by phytoplankton and excretion by Antarctic krill, Euphausia superba

In this study we examined the hypothesis that, under conditions of replete macronutrients and iron in the Southern Ocean, phytoplankton abundance and specific N uptake rates are influenced strongly by the processes of grazing and NH₄ regeneration. NH₄ and NO₃ uptake rates by marine phytoplankton were measured to the northeast and northwest of the island of South Georgia during January-February 1998. Mean specific uptake rate for NO₃ (vNO₃) was 0.0026 h⁻¹ (range 0.0013-0.0065 h⁻¹) and for NH₄ (vNH₄) was 0.0097 h⁻¹ (0.0014-0.0376 h⁻¹). vNH₄ was related positively with NH₄ availability, which ranged from 0.1 to 1.5 mmol m⁻³ within the upper mixed layer. Ambient NH₄ concentrations and vNH₄ were both positively related to local krill biomass values, computed from mean values along acoustic transect segments within 2 km of the uptake measurement stations. These biomass values ranged from ∼1 g krill fresh mass m⁻² in the northwest to >4 kg krill wet mass m⁻² in the northeast. In contrast to the variability found with NH₄ concentrations and uptake rates, vNO₃ was more uniform across the sampling sites. Under these conditions, increasing NH₄ concentration appeared to represent an additional N resource. However, high vNH₄ tended to be found for stations with lower phytoplankton standing stocks, across a total range of 0.24-20 mg chlorophyll a m⁻³. These patterns suggest a coupling between phytoplankton biomass, vNH₄ and krill in this region of variable but high krill biomass. Locally high concentrations of krill in parts of the study area appeared to have two opposing effects. On the one hand they could graze down phytoplankton stocks, but on the other hand, their NH₄ excretion supported enhanced uptake rates by the remaining, ungrazed cells.     

The planktonic food web of the Bizerte lagoon (south-western Mediterranean) during summer: I. Spatial distribution under different anthropogenic pressures

The structure and the trophic interactions of the planktonic food web were investigated during summer 2004 in a coastal lagoon of south-western Mediterranean Sea. Biomasses of planktonic components as well as bacterial and phytoplankton production and grazing by microzooplankton were quantified at four stations (MA, MB, MJ and R) inside the lagoon. Station MA was impacted by urban discharge, station MB was influenced by industrial activity, station MJ was located in a shellfish farming sector, while station R represented the lagoon central area. Biomasses and production rates of bacteria (7–33 mg C m⁻³; 17.5–35 mg C m⁻³ d⁻¹) and phytoplankton (80–299 mg C m⁻³; 34–210 mg C m⁻³ d⁻¹) showed high values at station MJ, where substantial concentrations of nutrients (NO₃⁻ and Si(OH)₄) were found. Microphytoplankton, which dominated the total algal biomass and production (>82%), were characterized by the proliferation of several chain-forming diatoms. Microzooplankton was mainly composed of dinoflagellates (Torodinium, Protoperidinium and Dinophysis) and aloricate (Lohmaniellea and Strombidium) and tintinnid (Tintinnopsis, Tintinnus, Favella and Eutintinnus) ciliates. Higher biomass of these protozoa (359 mg C m⁻³) was observed at station MB, where large tintinnids were encountered. Mesozooplankton mainly represented by Calanoida (Acartia, Temora, Calanus, Eucalanus, Paracalanus and Centropages) and Cyclopoida (Oithona) copepods, exhibited higher and lower biomasses at stations MA/MJ and MB, respectively. Bacterivory represented only 35% of bacterial production at stations MB and R, but higher fractions (65–70%) were observed at stations MA and MJ. Small heterotrophic flagellates and aloricate ciliates seemed to be the main controllers of bacteria. Pico- and nanophytoplankton represented a significant alternative carbon pool for micrograzers, which grazing represented 67–90% of pico- and nano-algal production in all stations. Microzooplankton has, however, a relatively low impact on microphytoplankton, as ≤45% of microalgal production was consumed in all stations. This implies that an important fraction of diatom production would be channelled by herbivorous meso-grazers to higher consumers at stations MA and MJ where copepods were numerous. Most of the microalgal production would, however, sink particularly at station MB where copepods were scare. These different trophic interactions suggest different food web structures between stations. A multivorous food web seemed to prevail in stations MJ and MA, whereas microbial web was dominant in the other stations.     

Influence of river discharge on plankton metabolic rates in the tropical monsoon driven Godavari estuary,India

To examine the influence of river discharge on plankton metabolic balance in a monsoon driven tropical estuary, daily variations in physico-chemical and nutrients characteristics were studied over a period of 15 months (September 2007 to November 2008) at a fixed location (Yanam) in the Godavari estuary, India. River discharge was at its peak during July to September with a sharp decrease in the middle of December and complete cessation thereafter. Significant amount of dissolved inorganic nitrogen (DIN, of 22–26 μmol l⁻¹) and dissolved inorganic phosphate (DIP, of 3–4 μmol l⁻¹) along with suspended materials (0.2–0.5 g l⁻¹) were found at the study region during the peak discharge period. A net heterotrophy with low gross primary production (GPP) occurred during the peak discharge period. The Chlorophyll a (Chl a) varied between 4 and 18 mg m⁻³ that reached maximum levels when river discharge and suspended loads decreased by >75% compared to that during peak period. High productivity was sustained for about one and half months during October to November when net community production (NCP) turned from net heterotrophy to autotrophy in the photic zone. Rapid decrease in nutrients (DIN and DIP by ∼15 and 1.4 μmol l⁻¹, respectively) was observed during the peak Chl a period of two weeks. Chl a in the post monsoon (October–November) was negatively related to river discharge. Another peak in Chl a in January to February was associated with higher nutrient concentrations and high DIN:DIP ratios suggest possible external supply of nitrogen into the system. The mean photic zone productivity to respiration ratio (P:R) was 2.38 ± 0.24 for the entire study period (September 2007–November 2008). Nevertheless, the ratio of GPP to the entire water column respiration was only 0.14 ± 0.02 revealing that primary production was not enough to support water column heterotrophic activity. The excess carbon demand by the heterotrophs could be met from the allochthonous inputs of mainly terrestrial origin. Assuming that the entire phytoplankton produced organic material was utilized, the additional terrestrial organic carbon supported the total bacterial activity (97–99%) during peak discharge period and 40–75% during dry period. Therefore, large amount of terrestrial organic carbon is getting decomposed in the Godavari estuarine system.     

Coloured dissolved organic matter (CDOM) in Southern North Sea waters: Optical characterization and possible origin

The variability and origin of the Coloured Dissolved Organic Matter (CDOM) were studied in the Belgian coastal and adjacent areas including offshore waters and the Scheldt estuary, through the parameters: absorption at 375 nm, a_CDOM(375), and the slope of the absorption curve, S. a_CDOM(375) varied between 0.20 and 1.31 m⁻¹ and between 0.97 and 4.30 m⁻¹ in the marine area and Scheldt estuary, respectively. S fluctuated between 0.0101 and 0.0203 nm⁻¹ in the marine area and between 0.0167 and 0.0191 nm⁻¹ in the Scheldt estuary. The comparative analysis of a_CDOM(375) and S variations evidenced different origins of CDOM in the BCZ. The Scheldt estuarine waters showed decreasing a_CDOM(375) values with increasing salinity but constant S value of ∼0.018 nm⁻¹ suggesting a dominant terrestrial origin of CDOM. On the contrary, samples collected in the marine domain showed a narrow range of a_CDOM(375) but highly variable S suggesting the additional presence of autochthonous sources of CDOM. This source was evidenced based on the sorting of the marine offshore data according to the stage of the phytoplankton bloom when they were collected. A clear distinction was made between CDOM released during the growth stage characterized by high S (∼0.017 nm⁻¹) and low a_CDOM(375) and the decay phase characterized by low S (∼0.013 nm⁻¹) and high a_CDOM(375). This observation was supported by CDOM measurements performed on pure phytoplankton cultures which showed increased CDOM release along the wax and wane of the bloom but decreasing S. We concluded that the high variability of the CDOM signature in offshore waters is explained by the local biological production and processing of CDOM.