DMSP and DMS dynamics and microzooplankton grazing in the Labrador Sea: application of the dilution technique

We adapted the dilution technique to study microzooplankton grazing of algal dimethylsulfoniopropionate (DMSP) vs. Chl a, and to estimate the impact of microzooplankton grazing on dimethyl sulfide (DMS) production in the Labrador Sea. Phytoplankton numbers were dominated by autotrophic nanoflagellates in the Labrador basin, but diatoms and colonial Phaeocystis pouchetii contributed significantly to phytomass at several high chlorophyll stations and on the Newfoundland and Greenland shelfs. Throughout the region, growth of algal Chl a and DMSP was generally high (0.2–1 d⁻¹), but grazing rates were lower and more variable, characteristic of the early spring bloom period. Production and consumption of Chl a vs. DMSP followed no clear pattern, and sometimes diverged greatly, likely because of their differing distributions among algal prey taxa and size class. In several experiments where Phaeocystis was abundant, we observed DMS production proportional to grazing rate, and we found clear evidence of DMS production by this haptophyte following physical stress such as sparging or filtration. It is possible that grazing-activated DMSP cleavage by Phaeocystis contributes to grazer deterrence: protozoa and copepods apparently avoided healthy colonies (as judged by relative growth and grazing rates of Chl a and DMSP), and grazing of Phaeocystis was significant only at one station where cells were in poor condition. Although we hoped to examine selective grazing on or against DMSP-containing algal prey, the dilution technique cannot differentiate selective ingestion and varying digestion rates of Chl a and DMSP. We also found that the dilution method alone was poorly suited for assessing the impact of grazing on dissolved sulfur pools, because of rapid microbial consumption and the artifactual release of DMSP and DMS during filtration. Measuring and understanding the many processes affecting organosulfur cycling by the microbial food web in natural populations remain a technical challenge that will likely require a combination of techniques to address.     

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, microzooplankton herbivory and community structure in the southeast Bering Sea: insight into the formation and temporal persistence of an Emiliania huxleyi bloom

Using the seawater dilution technique, we measured phytoplankton growth and microzooplankton grazing rates within and outside of the 1999 Bering Sea coccolithophorid bloom. We found that reduced microzooplankton grazing mortality is a key component in the formation and temporal persistence of the Emiliania huxleyi bloom that continues to proliferate in the southeast Bering Sea. Total chlorophyll a (Chl a) at the study sites ranged from 0.40 to 4.45 μg C l⁻¹. Highest phytoplankton biomass was found within the bloom, which was a mixed assemblage of diatoms and E. huxleyi. Here, 75% of the Chl a came from cells >10 μm and was attributed primarily to the high abundance of the diatom Nitzschia spp. Nutrient-enhanced total phytoplankton growth rates averaged 0.53 d⁻¹ across all experimental stations. Average growth rates for >10 μm and <10 μm cells were nearly equal, while microzooplankton grazing varied among stations and size fractions. Grazing on phytoplankton cells >10 μm ranged from 0.19 to 1.14 d⁻¹. Grazing on cells <10 μm ranged from 0.02 to 1.07 d⁻¹, and was significantly higher at non-bloom (avg. 0.71 d⁻¹) than at bloom (avg. 0.14 d⁻¹) stations. Averaged across all stations, grazing by microzooplankton accounted for 110% and 81% of phytoplankton growth for >10 and <10 μm cells, respectively. These findings contradict the paradigm that microzooplankton are constrained to diets of nanophytoplankton and strongly suggests that their grazing capability extends beyond boundaries assumed by size-based models. Dinoflagellates and oligotrich ciliates dominated the microzooplankton community. Estimates of abundance and biomass for microzooplankton >10 μm were higher than previously reported for the region, ranging from 22,000 to 227,430 cells l⁻¹ and 18 to 164 μg C l⁻¹. Highest abundance and biomass occurred in the bloom and corresponded with increased abundance of the large ciliate Laboea, and the heterotrophic dinoflagellates Protoperidinium and Gyrodinium spp. Despite low grazing rates on phytoplankton <10 μm within the bloom, the abundance and biomass of small microzooplankton (<20 μm) capable of grazing E. huxleyi was relatively high at bloom stations. This body of evidence, coupled with observed high grazing rates on large phytoplankton cells, suggests the phytoplankton community composition was strongly regulated by herbivorous activity of microzooplankton. Because grazing behavior deviated from size-based model predictions and was not proportional to microzooplankton biomass, alternate mechanisms that dictate levels of grazing activity were in effect in the southeastern Bering Sea. We hypothesize that these mechanisms included morphological or chemical signaling between phytoplankton and micrograzers, which led to selective grazing pressure.     

Egg production of Calanus finmarchicus—A basin-scale study

Egg production of Calanus finmarchicus was studied during joint basin-scale surveys in April–June 2003 in the Norwegian Sea. Surveys covered the whole Norwegian Sea and were conducted from Norwegian, Icelandic and Faroese research vessels. Stations were classified as being in pre-bloom, bloom or post-bloom phase according to levels of chlorophyll a and nitrate. Individual egg production rates and population egg production rates were calculated and compared between areas. Both individual egg production rates (eggs female⁻¹ day⁻¹) and population egg production rates (eggs m⁻² day⁻¹) were significantly higher in bloom areas compared with pre-bloom and post-bloom areas. However, when integrated over an estimated duration of the three phases, the time-integrated egg production (eggs m⁻²) in most years was highest in the pre-bloom phase, and this was explained by the longer duration of this phase compared with the two other phases.     

Reproductive success of Calanus pacificus during diatom blooms in Dabob Bay, Washington

While numerous laboratory studies in the last decade have shown that diatoms can induce reproductive failure in copepods, field evidence for a negative diatom effect is equivocal. To unambiguously elucidate the effects diatoms have on copepod reproduction in situ, we undertook a study of the abundance, distribution, grazing rates, and reproductive success of Calanus pacificus in Dabob Bay, Washington, USA, during two spring bloom periods. We simultaneously measured the phytoplankton composition, abundance, and distribution. Here we present results for the reproductive success of C. pacificus using four measurements: egg production rate, clutch size, egg hatching success, and naupliar survival (to the first feeding stage). Egg production rate was positively correlated with chlorophyll a concentration, and egg hatching success and naupliar survival were usually greater than 80%. However, in February 2002 and 2003 – during blooms of diatoms of the genus Thalassiosira – either egg hatching success, naupliar survival, or both were significantly depressed compared to other times in spring and summer. These effects, combined with evidence that C. pacificus was grazing aldehyde-producing Thalassiosira at the time of their blooms, indicate that diatoms can negatively affect copepod reproduction in the field, albeit only under specific circumstances and for brief periods.     

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.     

Factors controlling the timing of major spring bloom events in an UK south coast estuary

Factors controlling the timing of major (>10 mg chlorophyll a m⁻³) spring bloom events in the estuarine waters of the Solent, on the south coast of the UK, have been investigated. Winter to summer variations in chlorophyll a concentration together with relevant meteorological and hydrographical data have been analysed for 5 years (1988, 1992, 2001, 2002 and 2003). Mean water column irradiance is demonstrated to be the main factor controlling the timing of the first major spring bloom event, usually dominated by large chain-forming diatoms. When chlorophyll a concentration first exceeds 10 mg m⁻³ in spring (usually in May) the mean water column photosynthetic active radiation (PAR) averaged for one week prior to the sampling date was always >380 W h m⁻² d⁻¹. Prior to the main spring bloom event surface incident radiation and water turbidity combine to limit chlorophyll a concentration to levels <10 mg m⁻³. Chlorophyll a concentrations >10 mg m⁻³ do not occur in the Solent until almost the entire 10 m water column is within the euphotic zone (i.e. above 1% light level) and light extinction coefficient (k) is ca. ≤0.5 m⁻¹. Statistically, river flow explains the largest percentage of the variations in k and the delayed bloom in June 2002 is due to increased cloud cover and high levels of rainfall in May, which caused a reduction in surface incident irradiance and increased turbidity. Chlorophyll a peaks during these major bloom events generally occur on spring tides when increased mixing rates favour net growth of diatoms.     

Distribution and sinking rates of phytoplankton, detritus, and particulate biogenic silica in the Laptev Sea and Lena River (Arctic Siberia)

Concentrations and sinking rates of particulate biogenic silica (BSi), chlorophyll a (chl a) and phaeopigments (phae) (< 3 μm, 3–10 μm, > 10 μm and total), as well as the abundances of the major phytoplankton species, were studied during September 1991 in the Eastern Laptev Sea and the lower Lena River (Siberian Arctic). The highest chl a concentrations were found in two major “new” production regimes of the study area: (1) a deep chl a maximum (5.8 mg chl a m⁻³) (formed by the diatom Chaetoceros socialis) at 30 m depth on the outer shelf of the northern Laptev Sea, and (2) in the Lena River, where the phytoplankton community was dominated by fresh water diatoms (1.5 to 4.5 mg chl a m⁻³). Elevated chl a concentrations were also found in the river plume phytoplankton community (dominated by brackish water diatoms), NE of the Lena delta. In the Laptev Sea, the low chl a (0.1 to 3 mg chl a m⁻³) and high phae concentrations (0.5 to 14 mg phae m⁻³) indicated that the phytoplankton community (dominated by picoplanktic algae and nanoflagellates) was already senescent and affected by grazing losses. Biogenic silica values were highest in the Lena River (4 to 17 μM) as compared to the low values found in the Laptev Sea (0.3 to 4 μM). The large chl a size fraction, phae and BSi in the Lena River samples revealed the highest measured sinking rates (1.4, 2.3, and 1.5 m d⁻¹, respectively). The formation of a strong halocline, decreasing turbulence, and possible nutrient deficiency resulted in death, disintegration and rapid sedimentation of fresh water diatoms. This was accompanied by a decrease in the BSi concentration and growth of the picoplanktic size fraction (< 3 μm) in the estuarine mixing zone (Gulf of Buorkhaya). Only a minor part of BSi was bound to intact diatom cells (< 3%) in the surface layer, most of which being apparently associated with detrital particles. In the Lena River, approximately 12% of the total silica was bound to BSi fraction, yet elsewhere in the Laptev Sea and in the estuarine mixing zone the BSi:total silica ratio was ≤ 5%. Thus, the results reflected the successional stage of a late summer phytoplankton community, characterized by dominance of small autotrophs and patchy distribution of senescent diatoms no longer able to affect the relative high levels of dissolved silica supplied by the Lena River.     

Copepod grazing during spring blooms: Does Calanus pacificus avoid harmful diatoms?

During late winter and spring of 2002 and 2003, 24, 2–3 day cruises were conducted to Dabob Bay, Washington State, USA, to examine the grazing, egg production, and hatching success rates of adult female Calanus pacificus and Pseudocalanus newmani. The results of the copepod grazing experiments for C. pacificus are discussed here. Each week, copepod grazing incubation experiments from two different depth layers were conducted. Grazing was measured by both changes in chlorophyll concentration and cell counts. In 2002, there was one moderate bloom consisting mainly of Thalassiosira spp. in early February, and a larger bloom in April comprised of two Chaetoceros species and Phaeocystis sp. Similarly, in 2003, there were two blooms, an early one dominated by Thalassiosira spp., and a later one consisting of Chaetoceros spp. and Thalassiosira spp. Clearance rates on individual prey species, as calculated by cell counts, showed that C. pacificus are highly selective in their feeding, and may have much higher clearance rates on individual taxa than rates calculated from bulk chlorophyll disappearance. During weeks of high phytoplankton concentration, the copepods generally ate phytoplankton. However, they often rejected the most abundant phytoplankton species, particularly certain Thalassiosira spp., even though the rejected prey were often of the same genus and similar size to the preferred prey. It is speculated that this avoidance may be related to the possible deleterious effects that certain of these diatom species have on the reproductive success of these copepods. During weeks of medium to low phytoplankton concentration, the copepods selectively ate certain species of phytoplankton, and often had high electivity for microzooplankton. The selection mechanism must consist of active particle rejection most likely based on detection of surface chemical properties, since the diatoms that were selected were of the same genus, nearly the same size, and at lower numerical abundance than those cells that were avoided. The grazing choices made by these copepods may have important consequences for the overall ecosystem function within coastal and estuarine systems through changes in the transfer efficiency of energy to higher trophic levels.     

Seasonality of microzooplankton grazing in the northern Wadden Sea

A sequence of nine dilution experiments was conducted according to Landry and Hassett [Landry, M.R., Hassett, R.P., 1982. Estimating the grazing impact of marine microzooplankton. Mar. Biol. 67, 283–288] in the northern Wadden Sea from March until October 2004 to investigate the seasonality of microzooplankton grazing. From March until April, no grazing was observed. Microzooplankton grazing started in May (0.66 d^− 1) and increased until August (1.22 d^− 1). In October microzooplankton grazing was low again (0.17 d^− 1). Phytoplankton growth rates varied between 0 and 1.1 d^− 1. Since the reliability of dilution experiments is still frequently discussed in literature, we tested if our data obtained by dilution experiments reflected short-term in situ phytoplankton dynamics of the study site. We scaled experimental growth rates to water column irradiance, calculated short-term chlorophyll-a dynamics and compared the results to in situ measured chlorophyll-a concentrations. Calculated chlorophyll-a concentrations correlated significantly with in situ measured chlorophyll-a concentrations but slightly overestimated the in situ measured chlorophyll-a. This overestimation was in the range of phytoplankton assimilation reported for the Wadden Sea benthos. We will show that microzooplankton grazing had a large impact during the Phaeocystis bloom and during summer suggesting that a large proportion of phytoplankton biomass remained the pelagic food web. Microzooplankton grazing did not impact the diatom spring bloom and its demise.     

Temporal variation in phytoplankton composition related to water mass properties in the central part of Sagami Bay

Temporal variations in water mass properties and the composition of phytoplankton pigments in the central part of Sagami Bay were investigated by monthly observations from June 2002 to May 2004. Eleven pigments were quantified using high-performance liquid chromatography (HPLC) from 100%, 20%, and 5% light depths relative to the surface; the class-specific composition of phytoplankton community was then obtained by CHEMTAX analysis. The study area was influenced by the Kuroshio water for most of the observation period. The mean contribution of diatoms in all samples was relatively low (29%), while that of flagellates, mainly chlorophytes or cryptophytes, was quite high (60%). The phytoplankton composition at the three depths was uniform throughout the observation period, indicating that the vertical structure of the phytoplankton community did not develop significantly over time. A distinct temporal pattern was observed: flagellates dominated during the summer of 2002 and the winters of 2002–2003 and 2003–2004, while diatoms dominated during the summer of 2003. This pattern was associated with water mass changes. The community in the summer of 2003 was influenced by coastal water. While no distinct spring bloom of phytoplankton was observed, a weak increase in chlorophyll a was observed during the spring of 2004. Ocean color satellite data showed that fluctuations in chlorophyll a concentrations at time scales much shorter than a month occurred during the spring of 2003 and that the elevations in chlorophyll a levels were not continuous. The fluctuations were probably associated with rapid flushing by the Kuroshio water, which has low chlorophyll a content.     

黄海春季水华过程中微微型浮游生物的变化特点

于2007年3—4月在黄海中部海域采用流式细胞术研究了春季水华过程中聚球藻、微微型真核浮游生物和异养细菌的生物量变化。聚球藻和微微真核型浮游生物的生物量与叶绿素a浓度变化基本呈现相反的趋势,在水华前期较高,水华期迅速下降,直至水华后期又有所升高。异养细菌在整个水华过程中变化较小,生物量在水华期最高,与水柱叶绿素a浓度呈极显著正相关(r=0.319,p<0.01)。水华期这三类微微型浮游生物对浮游植物总碳生物量的贡献很低。纤毛虫和鞭毛虫捕食可能是导致聚球藻和微微型真核浮游生物在水华期生物量降低的主要原因。     

Seasonality of Plankton Assemblages in a Temperate Estuary

Abstract. Synoptic measurements of temperature, salinity, nutrients, primary productivity, chlorophyll a, and abundance and composition of phytoplankton, zooplankton, and ichthyoplank-ton were made over an annual cycle on the Peconic Bay estuary (Long Island, New York, USA). There were pronounced seasonal fluctuations in all variables measured. During the warmer season, the plankton was dominated by nanophytoplankton (athecate microflagellates and chlorophytes, short chains of diatoms), small zooplankton (copepod nauplii, copepodites, and adults of small copepod species) and gelatinous carnivores (ctenophores and medusae). During the colder season, the plankton was dominated (in terms of primary productivity and chlorophyll a) by netplankton, larger zooplankton (adult copepods) and fish larvae. The winter bloom of apparent netplankton (> 20 μm fraction) was largely an artifact of the screening method employed, in that long chains of a diatom with small individual cell size (Skeletonema costatum) comprised 84.4–97.8% of the phytoplankton present. There was a significant negative relationship over the year between length of diatom chains and number of smaller zooplankton. For this reason, as well as initiation of the winter bloom during a period of declining levels of both light and zooplankton, inception of the bloom appeared more related to release of zooplankton grazing pressure than to illumination. Temporally offset pulses of ctenophores and other zooplankton during the warmer season suggest substantial predation by ctenophores. Apparent decimation of copepod populations by ctenophore predation in late summer and fall immediately preceeded inception of the winter diatom bloom. Larval Ammodytes americanus were the dominant ichthyoplankton, and these co-occurred in winter with increased abundances of larger adult copepods of species upon which A. americanus is known to feed. With certain modifications the patterns recorded for Peconic Bay corresponded to both of two generalized trophic pathways proposed by Greve & Parsons (1977) for temperate waters: nanoplankton → small zooplankton → gelatinous zooplankton carnivores or netplankton → larger zooplankton → young fish. The former pattern characterized the warmer season, and the latter the colder season. Comparison of patterns in Peconic Bay with those in some other temperate estuarine and coastal waters suggests similarity, particularly for estuaries of the northeastern United States.     

The balance between microzooplankton grazing and phytoplankton growth in a highly productive estuarine fjord

During 24, three-day cruises to Dabob Bay, Washington State, USA, from February 4 to April 26, 2002, and February 4 to May 1 2003, we examined the relative growth and grazing rates of phytoplankton and microzooplankton using dilution experiments. Experiments were conducted over two time intervals: 8–10 h during the nighttime only, or 24 h from noon to noon. We used water from two depths during each cruise: from the surface mixed layer, and from a deep layer below the seasonal thermocline. During 2002, there was one mid-sized bloom consisting mainly of Thalassiosira spp. in early February, and a larger bloom in April comprised of two Chaetoceros spp. and Phaeocystis sp. During 2003, there were also two blooms, one in early February, which was again dominated by Thalassiosira spp., and a second larger bloom in mid-April, comprised mainly of Thalassiosira spp. and Chaetoceros spp. During all four of these blooms, and for both water source depths, specific grazing rates of microzooplankton were most often as high or higher than the calculated phytoplankton specific growth rates. The major microzooplankton categories that could have accounted for this were (1) a large Gyrodinium spp., (2) a group of fusiform-shaped mid-sized Protoperidinium species, and (3) three loosely defined taxonomic groups consisting of naked ciliates, tintinnids, and unidentified heterotrophic dinoflagellates. Based on our measurements, it appears that the microzooplankton community grazing pressure can often exert significant control on phytoplankton biomass, even during the extremely productive spring bloom periods and under several different diatom-dominated bloom types. These results suggest that even in highly productive estuarine ecosystems, which are often nurseries to economically important fisheries species, microzooplankton play a critical role and may significantly alter the availability and efficiency of transfer of energy to higher trophic levels.     

Inter-annual variability in phytoplankton summer blooms in the freshwater tidal reaches of the Schelde estuary (Belgium)

The inter-annual variability in phytoplankton summer blooms in the upper reaches of the Schelde estuary was investigated between 1996 and 2005 by monthly sampling at 10 stations. The large inter-annual variations of the chlorophyll a concentration in the freshwater tidal reaches were independent from variations in chlorophyll a in the tributary river Schelde. Summer mean chlorophyll a concentrations were significantly negatively correlated with flushing rate (Spearman correlation: r = −0.67, p = 0.05, n = 9) but not with temperature, irradiance and suspended particulate matter or dissolved silica (DSi) concentrations. During dry summers, low flushing rates permitted the development of dense phytoplankton populations in the upper part of the estuary, while during wet summers high flushing rates prevented the development of dense phytoplankton blooms. Flushing rate was also found to be important for the phytoplankton community composition. At low flushing rates, the community was dominated by diatoms that developed within the upper estuary. At high flushing rates, chlorophytes imported from the tributary river Schelde became more important in the phytoplankton community. The position of the chlorophyll a maximum shifted from the head of the estuary when flushing rates were low, to more downstream when flushing rates were high. Although DSi concentrations tended to be lower during years of high phytoplankton (mainly diatom) biomass, the relation with flushing rate was not significant.     

Ship-of-opportunity based phycocyanin fluorescence monitoring of the filamentous cyanobacteria bloom dynamics in the Baltic Sea

Distribution of cyanobacteria cannot be evaluated using chlorophyll a (Chla) in vivo fluorescence, as most of their Chla is located in non-fluorescing photosystem I. Phycobilin fluorescence, in turn, is noted as a useful tool in the detection of cyanobacterial blooms. We applied phycocyanin (PC) fluorometer in the monitoring of the filamentous cyanobacterial bloom in the Baltic Sea. For the bloom forming filamentous cyanobacteria Aphanizomenon flos-aquae and Nodularia spumigena, PC fluorescence maximum was identified using the excitation–emission fluorescence matrix. Consequently, the optical setup of our instrument was noted to be appropriate for the detection of PC, and with minor or no interference from Chla and phycoerythrin fluorescence, respectively.During summer 2005, the instrument was installed on a ferryboat commuting between Helsinki (Finland) and Travemünde (Germany), and data were collected during 32 transects providing altogether 200 000 fluorescence records. PC in vivo fluorescence was compared with Chla in vivo fluorescence and turbidity measured simultaneously, and with Chla concentration and biomass of the bloom forming filamentous cyanobacteria determined from discrete water samples.PC fluorescence showed a linear relation to the biomass of the bloom forming filamentous cyanobacteria, and the other sources of PC fluorescence are considered minor in the open Baltic Sea. Estimated by PC fluorescence, cyanobacterial bloom initiated late June at the Northern Baltic Proper, rapidly extended to the central Baltic Proper and the Gulf of Finland, and peaked in the mid-July with values up to 10 mg l⁻¹ (fresh weight). In late July, bloom vanished in most areas.During single transects, or for the whole summer, the variability in Chla concentrations was explained more by PC fluorescence than by Chla fluorescence. Thus, filamentous cyanobacteria dominated the overall variability in phytoplankton biomass. Consequently, we show that during the cyanobacterial blooms, the estimation of Chla concentration using only Chla in vivo fluorescence is not applicable, but PC in vivo fluorescence is required as a predictor as well.     

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.     

Mesoscale and microscale spatial variability of bacteria and viruses during a Phaeocystis globosa bloom in the Eastern English Channel

Sampling was conducted within inshore and offshore sites, characterized by highly dissimilar hydrodynamic and hydrobiological conditions, in the Eastern English Channel. The eutrophic inshore site was dominated by the influence of a dense bloom of the Prymnesiophyceae phytoplankton species Phaeocystis globosa, while the offshore site was characterized by more oceanic conditions. Within each site the microscale distributions of chlorophyll a and several flow cytometrically-defined subpopulations of heterotrophic bacteria and viruses were measured at a spatial resolution of 5 cm. The inshore site was characterized by comparatively high levels of microscale spatial variability, with concentrations of chlorophyll a, heterotrophic bacteria, and viruses varying by 8, 11 and 3.5-fold respectively across distances of several centimeters. Within the offshore site, microscale distributions of chlorophyll a and bacteria were markedly less variable than within the inshore site, although viruses exhibited slightly higher levels of heterogeneity. Significant mesoscale variability was also observed when mean microbial parameters were compared between the inshore and offshore sites. However, when the extent of change (max/min and coefficient of variation) was compared between meso- and microscales, the variability observed at the microscale, particularly in the inshore site, was substantially greater. This pattern suggests that microscale processes associated with Phaeocystis globosa bloom dynamics can generate heterogeneity amongst microbial communities to a greater degree than large scale oceanographic discontinuities.     

Evidence of microphytobenthic roles on coastal shallow water of the Seto Inland Sea,Japan

This study describes the temporal variation of microphytobenthic biomass and its controlling factors, as well as the impact of microphytobenthic activities on coastal shallow sediment in the eastern Seto Inland Sea, Japan. The chlorophyll a (Chl a), phaeopigments and sedimentary biophilic element (C, N, P and Si) contents in surface sediments, as well as nutrient concentrations at the sediment-water interface (overlying water and pore water) were observed monthly during November 2003 to May 2005 at one site in Shido Bay (water depth ca. 7 m) and at one site in Harima-Nada (35 m). No correlation was observed between the sedimentary biophilic elements and other parameters. The maximum chlorophyll a content of 48.2 μg g^–1 was found in surface sediments under the photon flux reaching the seafloor of 537 μmol photon m^–2 s^–1 during the winter period when water transparency was the highest at station S (Shido Bay). Our results suggest that higher chlorophyll a content in surface sediment was due to the fresh microphytobenthic biomass (mainly benthic diatom). We also found a significant negative correlation between Chl a and inorganic nutrients in pore water during the low temperature period, especially silicic acid. This result suggests that the silicic acid was assimilated largely during the increase of microphytobenthic biomass in surface sediment. This study suggests that the microphytobenthic roles may have a great effect on nutrient budgets during the large supply of irradiance (winter periods) for their photosynthetic growth in shallow coastal waters.     

Photosynthetic pigments as indicators of phytoplankton development during spring and summer in Adventfjorden (Spitsbergen)

The study was carried out from April 30 until July 13 of 1997 in Adventfjorden (Spitsbergen). Formation of less saline and warmer surface water (～1 m thick) caused by melting of the fast ice was observed in the fjord during the first days of May. In summer a less saline surface layer was about 3 m thick. Euphotic depth measured under ice sheet reached 12 m, whereas load of mineral matter brought with riverine discharge in summer (the content of total particulate matter in the fjord reached 1.66 kg m^?2) dramatically reduced euphotic zone depth to 0.35 m. By pigment measurement three phases of phytoplankton development in Adventfjorden were distinguished: (1) spring bloom that has started under fast ice and reached maximum in the mid of May, (2) stagnation period in June, (3) increase of pigment concentration in July, what could indicate a start of the next algae bloom. Analyses of chlorophylls and carotenoids revealed that diatoms (chl c, fucoxanthin), and green algae (chl b, lutein) dominated phytoplankton community in the fjord. Moreover, the presence of peridinin indicates the presence of Dinophyta and alloxanthin—the occurence of Cryptophyta. In May and June 1997 phytoplankton appeared mainly in the surface of water, while in July, as a result of inflow of turbulent riverine waters into Adventfjorden, algae cells were pushed down and the highest numbers were observed at the depth ～20 m. Great phaeopigments to chl a ratio (= 0.54) found in the fjord seston in June and July probably shows strong impact of zooplankton grazing on phytoplankton development. High contribution of chlorophyllide a in porphyrin a poll in samples collected under fast ice (chlorophyllide a/chl a ratio = 0.18) reflects the final stage of algal communitie succession in ice, just before spring ice melt and release of biota to oceanic water. Chloropyllide a content during summer was minor or not detectable, demonstrating that diatom cells were in good physiological condition. High chl a allomer/chl a ratio (average = 0.11 for the period investigated) confirms high oxygen concentration in environment of Adventfjorden.