Interannual variability in the spring phytoplankton bloom in Auke Bay,Alaska

Data on phytoplankton primary production, biomass, and species composition were collected during a 5 yr (1985–1989) study of Auke Bay, Alaska. The data were used to examine the interannual differences in the timing, duration, and magnitude of the spring phytoplankton blooms during each year and to relate these differences to interannual variations in weather patterns. Within any given year, a pre-bloom phase was characterized by low available light, low rates of primary production, low biomass, and predominantly small (<10µm) diatoms. During the primary bloom, integrated production rates rose to 4 to 4.5 g C m^–2 d^–1, and integrated biomass levels reached 415 to 972 mg chlorophyll m^–2. Primary blooms were usually dominated by large diatoms (Thalassiosira spp.), and in a single year (1989) byChaetoceros spp. The primary blooms terminated upon nutrient depletion in the euphotic zone. Secondary blooms, triggered by nutrient resupply from below, occurred sporadically after the primary bloom and accounted for 4 to 31% of total spring production. The date of initiation and the duration of the primary bloom varied little from year to year (standard deviation 3 and 5 d, respectively). Seasonal production rates and biomass levels varied interannually by a factor of 2 to 3. In contrast, intra-annual variations of more than an order of magnitude, especially in biomass, occurred over periods as short as 10 d. These large variations over short time periods indicate the importance of synchronous timing between spring blooms and the production of larval fish and shellfish, which depend on an appropriate and adequate food supply for growth and survival. Parameters describing primary production (e.g. peak daily production, mean daily production, and total production during the primary bloom and the entire season) exhibited little interannual variation (coefficient of variation, CV = 10 to 19%), but a large degree of intra-annual variation (CV = 77 to 116%). Similarly, interannual variations in biomass (peak chlorophyll, mean chlorophyll) were also lower (CV = 20 to 33%) than intra-annual variations (CV = 85 to 120%).     

Carbon budget for the spring bloom in Auke Bay,Alaska

This paper describes a carbon budget for the spring phytoplankton bloom in Auke Bay, a subarctic bay in southeastern Alaska. The budget was constructed using semiweekly data on carbon production, particulate carbon in the water column, and cumulative sedimentation of carbon, chlorophyll a, and pheopigments. From these measured parameters, seasonal carbon consumption, utilization, and import/export terms were derived. The chlorophyll and pheopigment data were used to partition carbon sinking out of the photic zone between phytoplankton cells and fecal material. The difference between total carbon production and carbon available for consumption was attributed primarily to carbon import/export related to advection of water masses into and out of the bay. Separate budgets were developed for each of five sampling years (1985–1989). An average of 130±16 g C/m² were produced by phytoplankton during each spring. Our model suggests that an average of 70% of this carbon was available for consumption by grazers within the bay; the remaining 30% is assumed to have been exported from the bay by advective transport. Of the available (non-exported) carbon, an average of 55% was consumed by grazers, 34% sank out of the photic zone in the form of uneaten algae, and about 11% remained at the end of the sampling period in the form of phytoplankton standing stocks. Overall, about 27% of the carbon produced each spring in Auke Bay (35 gC/m²) was used for growth and respiration by first-order consumers within the bay.     

Simulation model for wind-driven summer phytoplankton dynamics in Auke Bay,Alaska

A model was developed for use in simulating effects of short-term wind-mixing of the water column on estuarine phytoplankton dynamics. Simulation results, using parameter values estimated from field data, exhibited maxima observed in phytoplankton standing crop over a summer in Auke Bay, Alaska. Short-term wind-mixing of the water column can be important in determining the time of occurrence of phytoplankton community productivity pulses in deep estuaries.     

Occurrence of tri-n-butyltin-caused imposex in the North Pacific marine snail Nucella lima in Auke Bay,Alaska

Like female dog whelks (Nucella lapillus) in the Atlantic Ocean, females of the Pacific gastropod N. lima respond to low concentrations of tri-n-butyltin (TBT) by growing a penis and vas deferens. This condition, termed imposex, was found to occur in N. lima collected from August 1987 to May 1988 along a TBT pollution gradient associated with a marina in Auke Bay, Alaska. The suite of symptoms characteristic of imposex in N. lima was slightly different than in N. lapillus. Imposex, as measured by relative penis size (RPS) of females to males, increased from 0.0 to 34.27 along this gradient and as measured by vas deferens sequenceindex (VDS) increased similarly from 0.0 to 4.29. Concentrations of TBT in N. lima increased from below detection limits (about 0.010 g Sn g^-1 dry tissue wt) to 0.065 g Sn g^-1 dry tissue wt along the gradient. The gradient was determined by measuring TBT in whole-body tissues of bay mussels (Mytilus edulis). Concentrations of TBT in mussels increased from below detection limits to 0.833 g Sn g^-1 dry tissue wt in mussels from within the marina that was the major local source of TBT. Imposex, tissue TBT burden, and position along a TBT pollution gradient are significantly correlated (P<0.01) in N. lima. TBT was tested as a causative agent for imposex by exposing snails at a distant control site to TBT paint. After 1 mo exposure, 33% of the females grew a penis ranging in size from 0.2 to 0.8 mm. Our results generally corroborate those found for N. lapillus and indicate that imposex in N. lima is caused by environmental TBT exposure. We suggest that the RPS in the genus Nucella may be useful in monitoring TBT in coastal waters worldwide.     

Growth and collapse of a spring phytoplankton bloom in the Firth of Clyde,Scotland

A range of physical and chemical parameters were measured through the water column at two previously unstudied shallow-water sites in the Firth of Clyde, Scotland, in the spring of 1991. Sediment traps were also deployed at one of the sites. The results of this survey are presented and discussed. A bloom of Skeletonema costatum was observed at one site through March and into April, when it collapsed completely, the phytoplanktonic cells sedimenting out of the water column. No significant bloom or sedimentation was observed at the second study site.     

三峡水库香溪河库湾春季水华暴发藻类种源研究

2008年2月18日-4月17日,在距离香溪河河口约19 km处设一野外观测站,对观测站附近一定点进行持续监测,并在观测站进行围隔试验,探索香溪河库湾春季水华暴发藻类的种源,揭示水华暴发过程.监测结果表明:香溪河库湾春季硅藻(Diatom)(主要是针杆藻Synedra、星杆藻Asterionella)冰华藻类种源是原地水体,底泥对硅藻水华暴发影响不明显;2008年观测站附近的甲藻(Protoperidinium sp .)种源不是底泥中的孢囊.2008年春季香溪河库湾水华暴发过程可以分为"复苏-增长-衰亡"三个阶段,其中溶解性硅酸盐(D-Si)和光照是水华藻类复苏阶段的主要影响因子,而增长阶段的主要影响因子则是溶解性硅酸盐(D-Si)和总磷(TP),衰亡阶段由于4月8日骤降暴雨,浊度和降雨量是该阶段的主要影响因子.     

Egg production and feeding by copepods prior to the spring bloom of phytoplankton in Fram Strait,Greenland Sea

Reproductively activeCalanus hyperboreus (Krøyer) andC. glacialis Jaschnov were captured in the upper 100 m of Fram Strait (77° to 79°N) in late winter 1987. There was no evidence of a phytoplankton bloom; chlorophylla concentrations were uniformly low (<0.1 mg m^–3), and nitrate concentrations were uniformly high (>11.3 mg-at m^–3). Gut-fullness measurements indicated that females were ingesting very little. The maturation state of gonads of bothC. hyperboreus andC. glacialis indicated that 75% of females were in a ripe condition consistent with observed egg laying. The lipid content of females laying eggs was reduced in both species compared to that of females not laying eggs. InC. hyperboreus the reduction was 39% and inC. glacialis it was 44%. All the evidence suggests that bothC. hyperboreus andC. glacialis were laying eggs in late winter by using lipids stored previously; they were not relying on ambient concentrations of phytoplankton. The daily rate of egg laying byC. glacialis using lipids in late winter exceeded the rate reported for summer when ambient food supplies have been shown to be necessary. It is suggested that individuals, spawned well in advance of the spring bloom of phytoplankton, may comprise a major portion of the annual recruitment to the entire population ofC. glacialis in this area, and that their life cycle can be completed within 1 yr. NeitherMetridia longa (Lubbock) norC. finmarchicus (Gunnerus) laid eggs during this study.     

Copepod grazing during a declining spring phytoplankton bloom in the Northern North Sea

Grazing rates of larger (Calanus finmarchicus) and smaller (Acartia clausii Pseudocalanus elongatus etc.) copepods on naturally occurring phytoplankton populations were measured during a declining spring phytoplankton bloom. During the initial period, dominated by Chaetoceros spp. diatoms, constant ingestion rates were observed in Calanus finmarchicus at suspended particulate concentrations above 300 g carbon l^-1. Average daily intake during this time amounted to 35 to 40% of body carbon and reached a maximum of 50%. The feeding response of the smaller copepods was not so well defined, although a maximum daily intake of 56% body carbon was recorded. In both groups, feeding thresholds were at particulate concentrations around 50 g C l^-1. The feeding response of C. finmarchicus was correlated with both a change in their own population and in the food cell type. Linear regressions describing the concentration-dependent feeding response were: ingestion rate (IR)=1.16 total particulate volume (TPV)-36.15 during the initial part of the period compared with IR=0.41 TPV-12.18 for the latter period. C. finmarchicus filtered out slightly larger (x 1.2 diameter) particles than the small copepods and, in both groups, some filtering adjustment was made to accomodate to modal changes in the phytoplankton population from 20–30 m to 10 m diameter cells. Particle production during feeding was frequently evident in the smallest size ranges of particles and the ratio of particle production to ingestion rate was greater at low feeding rates.     

Further studies of short-term variation in the pigment composition of a spring phytoplankton bloom

Phytoplankton sampled at the chlorophyll maximum during the spring bloom at Loch Ewe, Scotland during 1986, were analysed using high-performance liquid chromatography for chloropigments and carotenoids. The day-to-day proportions of the pigments remained relatively stable throughout the bloom, with good preservation of chlorophyll a even at the sediment surface. Chlorophyllide a and phaeophorbide a appeared towards the end of the bloom. The major carotenoids remained stable throughout the bloom, but became rapidly degraded on sedimentation under the oxic conditions. The pigment data is related to the nutrient status of the bloom, when slow steady growth occurred without nutrient limitation. These data contrast sharply with data from the 1983 spring bloom at Loch Ewe when acute nutrient limitation took place.     

Sedimentation of particulate matter during a phytoplankton spring bloom in relation to the hydrographical regime

Data presented and discussed here were collected continuously during April/May 1975 in the Bornholm Basin of the Baltic Sea. Sedimentation rates of particulate matter were recorded with 5 multisample sediment traps from different depths in the water column at 2 positions 170 km apart. Current meter data collected during the same period and depths indicated that the positions remained hydrographically distinct during the investigation. Particulate matter from the euphotic zone including diatom cells formed the bulk of the material collected by all traps. This flux of organic particles to the bottom was unimpeded by the strong density stratification present in the water column. The upper traps always collected less material than lower ones. This paradox has been ascribed to diminishing current speeds with depth, concomitant with an increase in sinking rates of phytoplankton and phytodetritus. Both factors influence the sampling efficiency of sediment traps, which are thought to have underestimated actual sedimentation rates here. A time lag of 2 to 3 weeks in bloom development seemed responsible for the characteristic differences between the two positions. The phase of major sedimentation at one position covered about 18 days, and a distinct sequence in the composition of the material collected by the 6 glasses of each trap indicated phases of a progressively deteriorating phytoplankton population in the water column contributing the particulate material. A total of 6.2 g C m^-2 in 34 days was recorded at this station. Apart from a trap situated in an oxygen deficient layer which collected 0.44 g C m^-2 of zooplankton corpses, zooplankton mortality was overestimated by the traps. Large-scale sedimencation of fresh organic matter produced by the spring bloom is probably a regular feature in areas with low over-wintering zooplankton populations and, as such, possibly has a direct stimulatory effect on growth and reproduction of the benthos.Contribution No. 185 of the Joint Research Programme 95, Kiel University.     

Short-term variations in the pigment composition of a spring phytoplankton bloom from an enclosed experimental ecosystem

Phytoplankton samples taken during the spring bloom in the experimental enclosed ecosystem bags at Loch Ewe, Scotland, during 1983 were analysed for carotenoids and chlorophyll compounds using high-performance liquidchromatography (HPLC). Changes in the relative proportions of these pigments were related to day-to-day changes in the composition of the bloom and the physiological state of the algae. There is clear evidence for a change in the chlorophyllide a:chlorophyll a ratio, which reached a maximum as nutrient limitation occurred. No major qualitative changes in the carotenoid components were seen during the bloom; the relative proportion, however, of some carotenoids does provide useful information on the relative abundance of certain algal type in the phytoplankton.     

Benthic response to sedimentation of a spring phytoplankton bloom: Process and budget

The response of benthos to sedimentation of the spring phytoplankton bloom in the Kiel Bight (Western Baltic Sea) is described in terms of biomass (ATP) and activity (heat production and ETS-activity). Input of the bloom (11.5 g C m^-2) over a period from March 25 to April 19, 1980 to the sediment surface was in the form of cells and fresh phytodetritus as indicated by low C/N ratios (7) and high energy charge values (0.78). Benthic microbial activity was immediately stimulated by this input as heat production doubled and the activity of ETS tripled over winter values within 12 d in the absence of a significant increase in ambient temperature. A comparison of the two activity parameters suggests that anaerobic metabolism is more important during the winter (February and March) than after input of the bloom. Meiofauna was not able to take part in the first activity outburst. Benthic ATP-biomass (excluding macrofauna) doubled in late April due to microbial production, and doubled again in early May when meiofauna started reproductive activity. For macrofauna a general statement was not possible, although the sediment surface feeder Macoma baltica commenced a build up of glycogen and lipid resources immediately following bloom input whereas Nephtys ciliata, feeding on sediment and small macrofauna, showed a less pronounced and delayed effect from this input. An energy budget based on heat production measurements was calculated. A daily heat loss of the benthic community of 21.7 KJ m^-2 d^-1 (35.5 KJ m^-2 d^-1) was found, when a depth of 3 cm sediment (5 cm) was assumed. Heat production of macrofauna contributed less than 5% of this activity. The input of the bloom was burned within 21 (13) d. Preliminary estimations for an annual budget suggest that the vertical transport of particulate organic matter via sedimentation can only explain 25% (15%) of the benthic activity in the shallow water ecosystem of the Kiel Bight. This indicates the presence of other sources of organic carbon such as benthic primary production or other transport processes providing carbon to the sediments.Publication No. 384 of the Joint Research Program of Kiel University (Sonderforschungsbereich 95)     

The Baltic Sea spring phytoplankton bloom in a changing climate: an experimental approach

The response of the Baltic Sea spring bloom was studied in mesocosm experiments, where temperatures were elevated up to 6°C above the present-day sea surface temperature of the spring bloom season. Four of the seven experiments were carried out at different light levels (32–202?Wh?m^?2 at the start of the experiments) in the different experimental years. In one further experiment, the factors light and temperature were crossed, and in one experiment, the factors density of overwintering zooplankton and temperature were crossed. Overall, there was a slight temporal acceleration of the phytoplankton spring bloom, a decline of peak biomass and a decline of mean cell size with warming. The temperature influence on phytoplankton bloom timing, biomass and size structure was qualitatively highly robust across experiments. The dependence of timing, biomass, and size structure on initial conditions was tested by multiple regression analysis of the y-temperature regressions with the candidate independent variables initial light, initial phytoplankton biomass, initial microzooplankton biomass, and initial mesozooplankton (=copepod) biomass. The bloom timing predicted for mean temperatures (5.28°C) depended on light. The peak biomass showed a strong positive dependence on light and a weaker negative dependence on initial copepod density. Mean phytoplankton cell size predicted for the mean temperature responded positively to light and negatively to copepod density. The anticipated mismatch between phytoplankton supply and food demand by newly hatched copepod nauplii occurred only under the combination of low light and warm temperatures. The analysis presented here confirms earlier conclusions about temperature responses that are based on subsets of our experimental series. However, only the comprehensive analysis across all experiments highlights the importance of the factor light.     

Supply of organic matter to the sediment in the northern North Sea during a spring phytoplankton bloom

Strings of moored sediment traps were deployed in a 150 m water column over a period covering the growth and collapse of the spring bloom (4 April–3 June 1976) in an area of the northern North Sea. The efficiency of collection of material in the moored traps was compared to collections in free-drifting traps in the same area of deployment. The ways in which the data from the trap collections may be interpreted was considered at some length and a best estimate of the flux of organic carbon and nitrogen to the sediment was made. For the period prior to the spring bloom (4–23 April) this flux was 50 mg C m^–2 d^–1 (about 20% of primary production). During the bloom (24 April–19 May) it was about 185 mg C m^–2 d^–1 (35% of production) and during early summer (20 May–3 June) it was 115 mg C m^–2 d^–1, about 25% of the overlying production. The organic carbon and nitrogen content of the material collected was measured and the material was examined microscopically. There was evidence of a large settlement of diatoms immediately after the spring bloom which was reflected in changes in the C:N and C:chlorophyll ratios of the material collected. This change in biochemical composition of the material may affect its nutritional quality and have a stimulatory effect on the growth and reproduction of the animals living in the sediment.     

Vertical fluxes and ecological significance of organic materials during the phytoplankton bloom during austral summer in Breid Bay,Antarctica

A mooring system consisting of an in situ fluorometer at a depth of 50 m and a time-series sediment trap at a depth of 110 m was deployed at the sediment trap site (70°11.536'S; 24°18.679'E; water depth: 300 m) in Breid Bay, Antarctica in austral summer from 28 December 1985 to 13 February 1986. Sinking particles, consisting of diatoms (mainly Thalassiosira antarctica), were analyzed for organic materials, stable carbon and nitrogen isotopes. Vertical fluxes of organic carbon and nitrogen were determined to be within the ranges of 12.3 to 116 mg C m^-2 d^-1 and 1.79 to 15.4 mg N m^-2 d^-1, respectively, with maxima in the middle of January 1986, after which time the organic carbon and nitrogen fluxes tended to decrease with a steep gradient. High values of ¹³C were found in the organic matter of the sinking particles collected before the middle of January, indicating that the organic matter was derived from the diatoms in the logarithmic phase of their growth. Increased abundance of glucose was found in the water-extractable carbohydrate, which was one of the sinking particles collected in the middle of January. This fact clearly indicated that the diatoms were no longer in the growth phase but rather in the stationary phase, because reserved glucan as well as various cellular organic materials were reportedly accumulated within the algal cells in the stationary phase. Fifteen species of protein amino acids with trace amounts of -alanine, -aminobutyric acid and ornithine were found in the sinking particles upon acid hydrolysis, but the amino acid composition of these samples had not been affected much by biological agents. The essential amino acid index was calculated for the sinking particles collected in the course of the sediment trap experiment. The indices obtained indicated that the sinking particles collected in Breid Bay were more ecologically significant than the sinking and suspended particles from deep waters.     

Response of marine phytoplankton to nitrogen deficiency: Decreased nitrate uptake vs enhanced ammonium uptake

The uptake of nitrate and ammonium was measured separately in uni-algal, nitrogen-deficient cultures of four species of marine phytoplankton. Nitrogen-deficient phytoplankton took up ammonium at initial rates which greatly exceeded those measured for nitrogen-sufficient phytoplankton. However, nitrate uptake by nitrogendeficient cultures was generally much slower than either nitrate or ammonium uptake by nitrogen-sufficient cultures or ammonium uptake by nitrogen-deficient cultures. Considerable species differences were observed in the degree to which nitrogen deficiency increased ammonium uptake or decreased nitrate uptake. Loss of ability to take up nitrate, but enhanced ability to take up ammonium, as a result of nitrogen deficiency may be an adaptation to the different mechanisms by which nitrate and ammonium are supplied to the euphotic zone. In areas with an intermittent supply of nitrogen, changes in the ability of some species to take up nitrogen as a result of nitrogen starvation will influence species composition and complicate interpretations of measurements of nitrogen uptake.Contribution no. 1249 from the Department of Oceanography, University of Washington, and contribution no. 82006 from the Bigelow Laboratory for Ocean Sciences     

Analysis of phytoplankton pigments by HPLC before,during and after mass occurrence of the microflagellate Corymbellus aureus during the spring bloom in the open northern North Sea in 1983

Daily observations in April and May 1983 near a subsurface drifter launched in the Fladen Grounds area of the North Sea revealed that a large crop of the microflagellate Corymbellus aureus Green, producing up to 3 g C m^-2 d^-1, succeeded the diatoms dominating during the earlier phase of the spring bloom. The concentration of fucoxanthin was highest during the first half of May, while pigment fingerprints of high-performance liquid chromatograms of suspended matter sampled during the second half of May were dominated by 19-hexanoyloxyfucoxanthin, the main carotenoid of C. aureus (0.8x10^-9 mg cell^-1). This colonial Prymnesiophycean was described for the first time less than 10 years ago and has nerver been reported to be present in the North Sea. After 16 May the C. aureus population rapidly lost its growth potential, but the peak of the bloom in terms of cell number was found several days later (up to 9x10⁶ cells per dm³ on 19 May). The population then started to decline; senescence was associated with decreasing pigment contents of cells. The low growth rate of copepods registered during the second half of May was probably related to the poor quality of C. aureus as copepod food: the concentration of a phaeophorbide a typically found in copepod fecal pellets was highest during the diatom phase of the spring bloom preceding the C. aureus bloom.     

Uptake and release of phosphorus by phytoplankton in the Chesapeake Bay estuary,USA

The phytoplankton uptake and release rates for inorganic phosphate, dissolved organic phosphate and polyphosphate were estimated during 5 cruises on the Chesapeake Bay over a 9-month period. Phosphorus in all pools turned over in several minutes to 100 h, and each soluble pool appeared to contain fractions which were metabolically useful to the phytoplankton. Maximal uptake rates (V _m ) for orthophosphate ranged from 0.02 to 2.95 μg-at P (1.h)^-1 with half saturation constants (K _s ) between 0.09 and 1.72 μg-at P l^-1. At low soluble reactive phosphorus concentrations, the uptake rate of trace ³²P orthophosphate was initially rapid, but declined after 15 to 60 min incubation. The data suggest that the initial uptake phase was dominated by exchange of ³²PO₄ ^≡ for ³¹PO₄ ^≡ in the membrane transport systems whereas the subsequent phase represented the net incorporation of orthophosphate into phytoplankton cells.     

The spring phytoplankton bloom in Lindåspollene,a land-locked Norwegian fjord. II. Biomass and activity of net- and nanoplankton

From February 24 to April 24, weekly samples were collected at fixed depths at one station in Lindåspollene, a land-locked Norwegian fjord. Adenosine triphosphate (ATP), chlorophyll a, phaeophytin, ¹⁴C assimilation, and respiratory activity [electron transport system (ETS) activity] were measured in the net- (>30 m) and nanoplankton. Netplankton contained on the average 48% of the total chlorophyll a and 56% of the ATP, but contributed only 7% to the total carbon assimilation and 11% to the ETS activity. The assimilation numbers for net- and nanoplankton ranged from 0 to 1.2 and from 1.5 to 13.2, respectively. At the oxygen/hydrogen sulphide interface, high concentrations of ATP, but not of chlorophyll a, were found in the nanoplankton fraction. Netplankton algae grew actively only in the first phase of the bloom, and nanoplankton predominated later, apparently due to low nutrient concentrations. During the bloom, Skeletonema costatum made up the main part of the biomass. The number of cells in the chains decreased throughout the bloom, possibly reflecting the lowered silicate content. It appeared that only nanoplankton were grazed by zooplankton, while netplankton sank to the bottom and represented input to the benthos.