Phytoplankton blooms are strongly impacted by microzooplankton grazing in coastal North Pacific waters

Phytoplankton growth and microzooplankton grazing were measured in two productive coastal regions of the North Pacific: northern Puget Sound and the coastal Gulf of Alaska. Rates of phytoplankton growth (range: 0.09–2.69 day⁻¹) and microzooplankton grazing (range: 0.00–2.10 day⁻¹) varied seasonally, with lowest values in late fall and winter, and highest values in spring and summer. Chlorophyll concentrations also varied widely (0.19–13.65 μg l⁻¹). Large (>8 μm) phytoplankton cells consistently dominated phytoplankton communities under bloom conditions, contributing on average 65% of total chlorophyll biomass when chlorophyll exceeded 2 μg l⁻¹. Microzooplankton grazing was an important loss process affecting phytoplankton, with grazing rates equivalent to nearly two-thirds (64%) of growth rates on average. Both small and large phytoplankton cells were consumed, with the ratio of grazing to growth (g:μ) for the two size classes averaging 0.80 and 0.42, respectively. Perhaps surprisingly, the coupling between microzooplankton grazing and phytoplankton growth was tighter during phytoplankton blooms than during low biomass periods, with g:μ averaging 0.78 during blooms and 0.49 at other times. This tight coupling may be a result of the high potential growth and ingestion rates of protist grazers, some of which feed on bloom-forming diatoms and other large phytoplankton. Large ciliates and Gyrodinium-like dinoflagellates contributed substantially to microzooplankton biomass at diatom bloom stations in the Gulf of Alaska, and microzooplankton biomass overall was strongly correlated with >8 μm chlorophyll concentrations. Because grazing tended to be proportionally greater when phytoplankton biomass was high, the absolute amount of chlorophyll consumed by microzooplankton was often substantial. In nearly two-thirds of the experiments (14/23), more chlorophyll was ingested by microzooplankton than was available for all other biological and physical loss processes combined. Microzooplankton were important intermediaries in the transfer of primary production to higher trophic levels in these coastal marine food webs. Received: 12 November 1999 / Accepted: 4 October 2000     

Patterns of microzooplankton growth in dilution experiments across a trophic gradient: Implications for herbivory studies

To investigate the growth and grazing patterns of microzooplankton (MZP) in environments of differing productivity, dilution experiments measuring phytoplankton growth (μ) and grazing mortality (m) rates were performed using samples from contrasting locations along the Texas coast. Samples were collected from estuaries, coastal lagoons and offshore Gulf of Mexico locations in the spring and summer of 2001. MZP growth rates were determined in each dilution treatment. Although MZP biomass changed over time in most dilution treatments, adjusting μ and m for the actual grazer gradient (represented by geometric mean MZP biomass) did not cause a significant deviation from the nominal dilution gradient. Likewise, these adjustments did not yield significant regressions where none existed before adjustment. The dynamics of MZP taxonomic groups (ciliates, dinoflagellates) and size categories differed suggesting that in some cases internal predation may lead to trophic cascades. MZP biomass was higher in productive coastal waters and included a larger proportion of dinoflagellates than in the oligotrophic, ciliate-dominated waters of the Gulf of Mexico. The MZP biomass-to-chlorophyll a ratio was lowest in the hypereutrophic Nueces River, where MZP biomass significantly increased in all dilution treatments (net growth rates up to 2 day⁻¹) suggesting a strong top–down control. In the brown-tide dominated Upper Laguna Madre and the oligotrophic seagrass-dominated Lower Laguna Madre MZP growth was decoupled from that of phytoplankton. At these sites, MZP were likely fueled by bacterial carbon and mixotrophy, respectively. Observing the growth response of MZP in dilution experiments can provide insight into trophic structure and efficiency of the microbial food web.     

Feeding of Clausocalanids (Calanoida,Copepoda) on naturally occurring particles in the northern Gulf of Aqaba (Red Sea)

A total of 12 feeding experiments were conducted in the northern Gulf of Aqaba during spring (March/April) and autumn (September/October) 2002 at the Marine Science Station (MSS) in Aqaba. Females of three species of clausocalanids were selected: Clausocalanus farrani, C. furcatus and Ctenocalanus vanus. Natural occurring particle (NOP) larger than 5 μm were investigated as food source. The ambient chlorophyll a concentration at sampling depth (∼70 m) ranged between 0.15 and 1.00 μg chl a l⁻¹ and NOP concentrations ranged between 1.78 and 14.0 × 10³ cells l⁻¹ during the sampling periods. The division of particles into five size classes (5–10, 10–20, 20–50, 50–100 and >100 μm) revealed that most of the particles were found in the size classes below 50 μm (81–98%), while most of the natural occurring carbon (NOC) was concentrated in the size classes larger than 20 μm (70–95%). Ingestion rates were food density dependent rather than size dependent ranging between 0.02 and 1.65 × 10³ NOP ind⁻¹ day⁻¹ and 0.01 and 0.41 μg NOC ind⁻¹ day⁻¹, respectively, equivalent to a body carbon (BC) uptake between 0.4 and 51.8% BC day⁻¹. The share of the size classes to the total ingestion resembled in most cases the size class composition of the natural particle community.     

Life cycle of <Emphasis Type="Italic">Pseudocalanus acuspes</Emphasis> Giesbrecht (Copepoda,Calanoida) in the Central Baltic Sea: II. Reproduction,growth and secondary production

The population dynamics of Pseudocalanus acuspes in the Central Baltic Sea were studied from March 2002 to May 2003 on a monthly basis. All stages were present year round with a stage shift from nauplii to older copepodite stages over the course of the year. Biomass, estimated from prosome length, peaked between May and September with maximum recorded values of 594 and 855 mg C m⁻² in May 2002 and 2003, respectively. Differences in biomass between stations up to a factor of 20 were observed especially in April/May and October. Mean egg production rate (EPR) showed a seasonal course and was highest in April 2002 and 2003 with 3.6 and 2.1 eggs f⁻¹ day⁻¹, respectively, corresponding to a mean weight-specific egg production rate (SEPR) of 0.13 and 0.04. Egg production seems to be limited by food from May on. Stage durations determined from moulting experiments turned out to be extremely long. Maximum growth rates based on stage durations of 15–25 days at 4°C in May and July 2003 amounted for 0.03–0.05 day⁻¹ in CI-CIV. Comparing these rates with rates derived from temperature–development relationships for P. acuspes from the literature resulted in five times higher growth rates for the latter case. Secondary production reached values up to 9.1 mg C m⁻² day⁻¹ (method for continuously reproducing populations) and 10.5 mg C m⁻² day⁻¹ (increment summation).     

Exposure of the urban population to mercury in Changchun city, Northeast China

An assessment of exposure to mercury in Changchun city has been undertaken. We estimated Hg exposure to members of the general population based on currently available information and our research. We also studied the Hg concentrations in scalp hair of adults. Adults have an estimated intake of all Hg species via all routes of 6.780 μg day⁻¹ (excluding dental amalgam), which equates to an absorbed dose of 1.718 μg day⁻¹. Fish consumption was the most important exposure route (12% of intake, 43% of absorbed dose). Furthermore, air, cereals and vegetables were important exposure routes, and these exposure were estimated for absorbed dosed at 0.296, 0.209 and 0.318 μg day⁻¹, respectively. The mean Hg concentration in hair was 0.448 μg g⁻¹ (range 0.092–10.463 μg g⁻¹). Hg concentration in the hair of males was 0.422 μg g⁻¹ (0.105–2.665 μg g⁻¹), and was 0.474 μg g⁻¹(0.092–10.463 μg g⁻1) in the hair of females. Neither place of residence nor age had any significant effect on hair Hg concentrations.     

Microbial activity and carbon, nitrogen, and phosphorus content in a subtropical seagrass estuary (Florida Bay): evidence for limited bacterial use of seagrass production

Bacterial abundance, production, and extracellular enzyme activity were determined in the shallow water column, in the epiphytic community of Thalassia testudinum, and at the sediment surface along with total carbon, nitrogen, and phosphorus in Florida Bay, a subtropical seagrass estuary. Data were statistically reduced by principle components analysis (PCA) and multidimensional scaling and related to T. testudinum leaf total phosphorus content and phytoplankton biomass. Each zone (i.e., pelagic, epiphytic, and surface sediment community) was significantly dissimilar to each other (Global R = 0.65). Pelagic aminopeptidase and sum of carbon hydrolytic enzyme (esterase, peptidase, and α- and β-glucosidase) activities ranged from 8 to 284 mg N m⁻² day⁻¹ and 113–1,671 mg C m⁻² day⁻¹, respectively, and were 1–3 orders of magnitude higher than epiphytic and sediment surface activities. Due to the phosphorus-limited nature of Florida Bay, alkaline phosphatase activity was similar between pelagic (51–710 mg P m⁻² day⁻¹) and sediment (77–224 mg P m⁻² day⁻¹) zones but lower in the epiphytes (1.1–5.2 mg P m⁻² day⁻¹). Total (and/or organic) C (111–311 g C m⁻²), N (9.4–27.2 g N m⁻²), and P (212–1,623 mg P m⁻²) content were the highest in the sediment surface and typically the lowest in the seagrass epiphytes, ranging from 0.6 to 8.7 g C m⁻², 0.02–0.99 g N m⁻², and 0.5–43.5 mg P m⁻². Unlike nutrient content and enzyme activities, bacterial production was highest in the epiphytes (8.0–235.1 mg C m⁻² day⁻¹) and sediment surface (11.5–233.2 mg C m⁻² day⁻¹) and low in the water column (1.6–85.6 mg C m⁻² day⁻¹). At an assumed 50% bacterial growth efficiency, for example, extracellular enzyme hydrolysis could supply 1.8 and 69% of epiphytic and sediment bacteria carbon demand, respectively, while pelagic bacteria could fulfill their carbon demand completely by enzyme-hydrolyzable organic matter. Similarly, previously measured T. testudinum extracellular photosynthetic carbon exudation rates could not satisfy epiphytic and sediment surface bacterial carbon demand, suggesting that epiphytic algae and microphytobenthos might provide usable substrates to support high benthic bacterial production rates. PCA revealed that T. testudinum nutrient content was related positively to epiphytic nutrient content and carbon hydrolase activity in the sediment, but unrelated to pelagic variables. Phytoplankton biomass correlated positively with all pelagic components and sediment aminopeptidase activity but negatively with epiphytic alkaline phosphatase activity. In conclusion, seagrass production and nutrient content was unrelated to pelagic bacteria activity, but did influence extracellular enzyme hydrolysis at the sediment surface and in the epiphytes. This study suggests that seagrass-derived organic matter is of secondary importance in Florida Bay and that bacteria rely primarily on algal/cyanobacteria production. Pelagic bacteria seem coupled to phytoplankton, while the benthic community appears supported by epiphytic and/or microphytobenthos production.     

Reproduction in Balanus amphitrite Darwin (Cirripedia: Thoracica): influence of temperature and food concentration

Balanus amphitrite, an acorn barnacle, is distinctly euryhaline, eurythermal and a dominant fouling organism found in warm and temperate waters throughout the world. In this study, the influence of temperature and food concentration on the reproductive biology of this species collected from a tropical habitat was evaluated. Adult barnacles were maintained at 20, 25 and 30°C temperatures at different concentrations of food (50, 100, 150 and 200 Artemia ind⁻¹ day⁻¹). In this previously believed obligatory cross-fertilizing hermaphrodite, self-fertilization was observed. The rise in temperature from 20 to 30°C resulted in a longer interbreeding interval (6–7 days, 200 Artemia ind⁻¹ day⁻¹; 11–13 days, 50 Artemia ind⁻¹ day⁻¹). Computed carbon gained through feeding during the interbreeding interval indicated an inverse relationship to the temperature. At 20°C, although a greater amount of carbon was gained through feeding, the numbers of larvae produced were fivefold less when compared to those raised at 30°C. At 20°C, 2.3 μg C was required to produce a single larva, whereas at 30°C it was 0.4 μg C. A rise in rearing temperature also influenced the molting rate positively. Observations on temporal variation in the gonad development of this species in a tropical coastal environment influenced by the monsoons indicated gonad development to be positively related to chlorophyll a concentration.     

Spring sea ice photosynthesis,primary productivity and biomass distribution in eastern Antarctica, 2002–2004

While it is known that Antarctic sea ice biomass and productivity are highly variable over small spatial and temporal scales, there have been very few measurements from eastern Antarctic. Here we attempt to quantify the biomass and productivity and relate patterns of variability to sea ice latitude ice thickness and vertical distribution. Sea ice algal biomass in spring in 2002, 2003 and 2004 was low, in the range 0.01–8.41 mg Chl a m⁻², with a mean and standard deviation of 2.08 ± 1.74 mg Chl a m⁻² (n = 199). An increased concentration of algae at the bottom of the ice was most pronounced in thicker ice. There was little evidence to suggest that there was a gradient of biomass distribution with latitude. Maximum in situ production in 2002 was approximately 2.6 mg C m⁻² h⁻¹ with assimilation numbers of 0.73 mg C (mg Chl a)⁻¹ h⁻¹. Assimilation numbers determined by the ¹⁴C incubations in 2002 varied between 0.031 and 0.457 mg C (mg Chl a)⁻¹ h⁻¹. Maximum fluorescence quantum yields of the incubated ice samples in 2002 were 0.470 ± 0.041 with E _k indices between 19 and 44 μmol photons m⁻² s⁻¹. These findings are consistent with the shade-adapted character of ice algal communities. In 2004 maximum in situ production was 5.9 mg C m⁻² h⁻¹ with an assimilation number of 5.4 mg C (mg Chl a)⁻¹ h⁻¹. Sea ice biomass increased with ice thickness but showed no correlation with latitude or the time the ice was collected. Forty-four percent of the biomass was located in bottom communities and these were more commonly found in thicker ice. Surface communities were uncommon.     

The tunicate Salpa thompsoni ecology in the Southern Ocean. I. Distribution, biomass, demography and feeding ecophysiology

Distribution, density, and feeding dynamics of the pelagic tunicate Salpa thompsoni have been investigated during the expedition ANTARKTIS XVIII/5b to the Eastern Bellingshausen Sea on board RV Polarstern in April 2001. This expedition was the German contribution to the field campaign of the Southern Ocean Global Ocean Ecosystems Dynamics Study (SO-GLOBEC). Salps were found at 31% of all RMT-8 and Bongo stations. Their densities in the RMT-8 samples were low and did not exceed 4.8 ind m⁻² and 7.4 mg C m⁻². However, maximum salp densities sampled with the Bongo net reached 56 ind m⁻² and 341 mg C m⁻². A bimodal salp length frequency distribution was recorded over the shelf, and suggested two recent budding events. This was also confirmed by the developmental stage composition of solitary forms. Ingestion rates of aggregate forms increased from 2.8 to 13.9 μg (pig) ind⁻¹ day⁻¹ or from 0.25 to 2.38 mg C ind⁻¹ day⁻¹ in salps from 10 to 40 mm oral-atrial length, accounting for 25–75% of body carbon per day. Faecal pellet production rates were on average 0.08 pellet ind⁻¹ h⁻¹ with a pronounced diel pattern. Daily individual egestion rates in 13 and 30 mm aggregates ranged from 0.6 to 4.8 μg (pig) day⁻¹ or from 164 to 239 μg C day⁻¹. Assimilation efficiency ranged from 73 to 90% and from 65 to 76% in 13 and 30 mm aggregates, respectively. S. thompsoni exhibited similar ingestion and egestion rates previously estimated for low Antarctic (~50°S) habitats. It has been suggested that the salp population was able to develop in the Eastern Bellingshausen Sea due to an intrusion into the area of the warm Upper Circumpolar Deep Water     

Functional responses of copepod nauplii using a high efficiency gut fluorescence technique

To investigate copepod nauplii ingestion rates on phytoplankton, we have adapted the traditional gut fluorescence technique as it can be used with lower gut pigment concentrations. With the improved technique, laboratory experiments were performed to estimate functional responses for nauplii of Calanus helgolandicus and Centropages typicus. Nauplii were raised from eggs to copepodites and the experiments were performed with stages NIV-NV. Gut evacuation rates and ingestion rates were measured on Isochrysis galbana at different concentrations. Specific ingestion rates ranged between 0.038–0.244 μg C μg⁻¹ nauplii C d⁻¹ for C. typicus and 0.041–1.412 μg C μg⁻¹ nauplii C d⁻¹ for C. helgolandicus. Both species showed a type III functional response, reaching a saturation concentration at around 600 μgC l⁻¹ for C. typicus and 800 μgC l⁻¹ for C. helgolandicus. An erratum to this article can be found at     

Growth and development rates of Calanus finmarchicus nauplii during a diatom spring bloom

Growth and development rates were determined for nauplii of Calanus finmarchicus (Gunnerus) in the near-shore waters of a western Norwegian fjord from in situ mesocosm incubations. The major food source for the nauplii was diatoms, but Phaeocystis sp., dinoflagellates and ciliates were also part of the diet. At local temperatures ranging from 4.8 to 5.2 °C the cumulative median development time from hatching to Nauplius VI was 19 d. The time taken to molt to the next naupliar stage was approximately constant (3 d) from Stages IV to VI, but Stage III needed the longest development time (5 d). The instantaneous growth rate in terms of body carbon was negative from hatching to Nauplius Stage II, but as high as 0.25 to 0.30 d⁻¹ from Stage III to V. Enhancement of food resources by nutrient addition led to no significant change in specific growth rates. Additionally, the cohorts from different nutrient regimes showed almost equal development time, size and body carbon within stages. Length–weight relationships of nauplii from the two different food resources were: W _{low resources} = 4.17 × 10⁻⁶ × L ^2.03 (r ² = 0.84) and W _{high resources} = 4.29 × 10⁻⁶ × L ^2.05 (r ² = 0.92), where weight (W) is in micrograms of C and body length (L) in micrometers. The natural body morphology of naupliar stages I to VI is illustrated with digital images, including the final molt from Nauplius VI to Copepodid Stage I. In general, development of the nauplii was faster than that of the copepodids of C. finmarchicus, and structural growth was exponential from naupliar stages III to VI. This study validates our earlier results that nauplii of C. finmarchicus can obtain high growth and nearly maximal developmental rates at relatively low food levels (∼50 μg C l⁻¹), suggesting that nauplii exhibit far less dependence on food supply than copepodids. Received: 30 July 1999 / Accepted: 7 March 2000     

Habitat use,urea production and spawning in the gulf toadfish <Emphasis Type="Italic">Opsanus beta</Emphasis>

Field studies were conducted in Johnson Key Basin, Florida Bay, USA from September 2002 through September 2004 to examine physiological, ecological, and behavioral characteristics of the gulf toadfish, Opsanus beta (Goode and Bean in Proc US Natl Mes 3:333–345, 1880), in relation to nitrogen metabolism, habitat usage, and spawning. Fish collected 5 cm above sediments in experimental shelters (epibenthic) were compared with those collected by throw traps which were found on or burrowing within sediments. The relationship between microhabitat ammonia and urea excretion, as determined by the enzymatic activity of glutamine synthetase (GS), was examined. The hypothesis tested was that O. beta occupying epibenthic nests were less ureotelic with lower GS activities than non-nesting individuals on/in sediments, due to a decreased environmental ammonia burden. Porewater total ammonia (T _Amm) concentrations at a sediment depth of 5 cm, i.e., the approximate depth of burrowing toadfish, ranged from 0 to 106.5 μmol N l⁻¹ while the pH ranged from 7.48 to 9.14. There was a weak but significant correlation between environmental ammonia (NH₃) concentration and hepatic GS activity for epibenthic toadfish (P < 0.001, r ² = 0.10), but not for burrowing toadfish. Mean urea-N and T _Amm concentrations within shelters occupied by toadfish (n = 281) were 9.8 ± 0.83 μmol N l⁻¹ and 13.0 ± 0.7 μmol N l⁻¹, respectively. As predicted, hepatic GS activity was significantly lower in epibenthic toadfish captured in shelters (4.40 ± 0.24 μmol min⁻¹ g⁻¹; n = 281) as compared to individuals on/in sediments (6.61 ± 0.47 μmol min⁻¹ g⁻¹; n = 128). Glutamine synthetase activity generally peaked in March (spawning season) and was lowest in July. Gender differences in hepatic and branchial GS activity were also found during the spawning season, which is attributable to the fact that males brood and guard offspring in their epibenthic nests while females often rest on or burrow into the sediments. Finally, hepatic and branchial GS appeared to have different patterns of enzymatic activity suggesting functional differences in gene expression.     

Phytoplankton community structure and primary production in small intertidal estuarine-bay ecosystem (eastern English Channel,France)

From May 2002 to October 2003, a fortnightly sampling programme was conducted in a restricted macrotidal ecosystem in the English Channel, the Baie des Veys (France). Three sets of data were obtained: (1) physico-chemical parameters, (2) phytoplankton community structure illustrated by species composition, biovolume and diversity, and (3) primary production and photosynthetic parameters via P versus E curves. The aim of this study was to investigate the temporal variations of primary production and photosynthetic parameters in this bay and to highlight the potential links with phytoplankton community structure. The highest level of daily depth-integrated primary production Pz (0.02–1.43 g C m⁻² d⁻¹) and the highest maximum photosynthetic rate P ^B _max (0.39–8.48 mg C mg chl a ⁻¹ h⁻¹) and maximum light utilization coefficient α^B [0.002–0.119 mg C mg chl a ⁻¹ h⁻¹ (μmol photons m⁻² s⁻¹)] were measured from July to September. Species succession was determined based on biomass data obtained from cell density and biovolume measurements. The bay was dominated by 11 diatoms throughout the year. However, a Phaeocystis globosa bloom (up to 25 mg chl a m⁻³, 2.5 × 10⁶ cells l⁻¹) was observed each year during the spring diatom bloom, but timing and intensity varied interannually. Annual variation of primary production was due to nutrient limitation, light climate and water temperature. The seasonal pattern of microalgal succession, with regular changes in composition, biovolume and diversity, influenced the physico-chemical and biological characteristics of the environment (especially nutrient stocks in the bay) and thus primary production. Consequently, investigation of phytoplankton community structure is important for developing the understanding of ecosystem functioning, as it plays a major role in the dynamics of primary production.     

Strong variability in bacterioplankton abundance and production in central and western Bay of Bengal

With large influx of freshwater that decreases sea-surface salinities, weak wind forcing of <10 m s⁻¹ and almost always warm (>28°C) sea-surface temperature that stratifies and shallows the mixed layer leading to low or no nutrient injections into the surface, primary production in Bay of Bengal is reportedly low. As a consequence, the Bay of Bengal is considered as a region of low biological productivity. Along with many biological parameters, bacterioplankton abundance and production were measured in the Bay of Bengal during post monsoon (September–October 2002) along an open ocean transect, in the central Bay (CB, 88°E) and the other transect in the western Bay (WB). The latter representing the coastal influenced shelf/slope waters. Bacterioplankton abundances (<2 × 10⁹cells l⁻¹) were similar to those reported from the HNLC equatorial Pacific and the highly productive northern Arabian Sea. Yet, the thymidine uptake rates along CB (average of 1.46 pM h⁻¹) and WB (average of 1.40 pM h⁻¹) were less than those from the northwestern Indian Ocean. These abundances and uptake rates were higher than those in the oligotrophic northwestern Sargasso Sea (<7 × 10⁸ cells l⁻¹; av 1.0 pM h⁻¹). Concentrations of chlorophyll a (chl a), primary production rates and total organic carbon (TOC) were also measured for a comparison of heterotrophic and autotrophic production. In the WB, bacterioplankton carbon biomass equaled ∼ 95% of chl a carbon than just 31% in the CB. Average bacterial:primary production (BP:PP) ratios accounted for 29% in the CB and 31% in the WB. This is mainly due to lower primary productivity (PP) in the WB (281 mg C m⁻² d⁻¹) than in the CB (306 mg C m⁻² day⁻¹). This study indicates that bacteria–phytoplankton relationship differs in the open (CB) and coastal waters (WB). Higher abundance and contrastingly low bacterial production (BP) in WB may be because of the riverine bacteria, brought in through discharges, becoming dormant and unable to reproduce in salinities of 28 or more psu. Heterotrophic bacteria appear to utilize in situ DOC rather rapidly and their carbon demand is ∼50% of daily primary production. It is also apparent that allochthonous organic matter, in particular in the western Bay, is important for meeting their carbon demand.     

Experimental evidence for internal predation in microzooplankton communities

The fate of microzooplankton production, whether it is channeled to mesozooplankton or recycled within the microbial food web, has major implications for the oceanic carbon cycle. The aim of this study was to estimate internal predation within naturally occurring microzooplankton communities. A dilution series based on the Landry and Hasset technique was created by mixing 200-μm-screened water (used as whole water) with 5-μm-screened seawater due to the dominance of pico- and small nanoplankton at our study site. This modification of the original technique allows for gradual reduction in microzooplankton abundance and thus internal predation while maintaining sufficient phytoplankton prey levels for microzooplankton growth in diluted treatments. Microzooplankton growth and mortality rates were calculated based on the changes in abundance during 24-h incubation. In the diluted treatments, microzooplankton growth rates were significantly higher (1.21 ± 0.20 day^?1 for ciliates and 0.88 ± 0.05 day^?1 for heterotrophic dinoflagellates) compared to those in whole seawater where microzooplankton abundance remained unchanged or even declined over time. Approximately 79 % of microzooplankton production was consumed within the microzooplankton, with aloricate ciliates being the most vulnerable to predation. These findings support the assumption that trophic interactions between microzooplankton can be an important factor controlling their production and, thus, energy transfer in picoplankton-dominated pelagic ecosystems.     

Metabolism and elemental composition of four oncaeid copepods in the western subarctic Pacific

Respiration rates and elemental composition (carbon and nitrogen) were determined for four dominant oncaeid copepods (Triconia borealis, Triconia canadensis, Oncaea grossa and Oncaea parila) from 0–1,000 m depth in the western subarctic Pacific. Across the four species of which dry weight (DW) varied from 2.0 to 32 μg, respiration rates measured at in situ temperature (3°C) increased with DW, ranging from 0.84 to 7.4 nl O₂ individual⁻¹ h⁻¹. Carbon (C) and nitrogen (N) composition of the four oncaeid species ranged from 49–57% of DW and 7.0–10.3% of DW, respectively, and the resultant C:N ratios were 4.8–8.3. The high C contents and C:N ratios were reflected by large accumulation of lipids in their body. Specific respiration rates (SR, a fraction of body C respired per day) ranged between 0.5 and 1.3% day⁻¹. Respiration rates adjusted to a body size of 1 mg body N (i.e. adjusted metabolic rates, AMR) of the four oncaeid species [0.6–1.1 μl O₂ (mg body N)^−0.8 h⁻¹ at 3°C] were significantly lower than those (1.7–5.1) reported in the literature for oithonid and calanoid copepods at the same temperature. The present results indicate that lower metabolic expenditure due to less active swimming (pseudopelagic life mode) together with rich energy reserve in the body (as lipids) are the characters of oncaeid copepods inhabiting in the epi- and mesopelagic zones of this region.     

Standing crops and vertical distribution of four groups of marine planktonic ciliates in relation to phytoplankton chlorophyll a

Standing crops and the vertical distribution of four groups of ciliates, autotrophic naked ciliates (ANC), heterotrophic naked ciliates (HNC ), mixotrophic naked ciliates (MNC) and loricated ciliates (LC ), were analysed in relation to phytoplankton chlorophyll a in the western Pacific. Data were gathered from the upper 60 to 200 m of water in the subarctic North Pacific in spring, in the subtropical North Pacific in spring, in Toyama Bay in summer, off eastern Australia in spring and off Sanriku in spring and fall. Of the four groups, the standing crop of HNC (cells l⁻¹) showed the highest correlation to phytoplankton chloro- phyll a [CHL, μg l⁻¹; r=0.66, n=365 (HNC=490 ×CHL ^0.653)]. Depths of the maximum layers of HNC and MNC were usually shallower than that of chlorophyll a, while those of ANC and LC occurred frequently beneath the chlorophyll maximum layer. This indicates that these four ciliate groups are substantially different from each other, and that each group holds a different ecological position in the marine microbial food web. Received: 27 October 1997 / Accepted: 27 June 1998     

Response of the microbial food web to manipulation of nutrients and grazers in the oligotrophic Gulf of Aqaba and northern Red Sea

The control mechanisms within the pelagic microbial food web of the oligotrophic Gulf of Aqaba and the northern Red Sea were investigated in the spring of 1999. Nutrient conditions and potential grazer impact were manipulated in a series of dilution experiments. Ambient nutrient concentrations and autotrophic biomass were very low (0.23–1.21 µmol NO₃ l^–1, 0.06–0.98 µmol NH₄ l^–1, 1.08–1.17 µmol Si l^–1, 0.08–0.12 µmol P l^–1, 0.15–0.36 µg chlorophyll a l^–1). The planktonic community was characterized by low abundances [3.0–5.5×10⁵ heterotrophic bacteria ml^–1, 0.58–7.2×10³ ultraphytoplankton <8 µm ml^–1 (small eukaryotic photoautotrophs and Prochlorococcus sp., excluding Synechococcus sp.), 0.45–4.4×10⁴ Synechococcus sp. ml^–1, 0.32–1.2×10³ heterotrophic nanoflagellates ml^–1, 1.3–3.8×10³ phytoplankton >8 µm l^–1, 0.93–5.4×10² microzooplankton l^–1] and dominated by small forms (0.2–8 µm). Dinoflagellates and oligotrichous ciliates were the most common groups in initial samples among the phytoplankton >8 µm and microzooplankton, respectively. Results show that bottom-up and top-down control mechanisms operated simultaneously. Small organisms were vulnerable to grazing, with maximum grazing rates of 1.1 day^–1 on heterotrophic bacteria and 1.3 day^–1 on ultraphytoplankton. In contrast, algae >8 µm showed stronger signs of nutrient limitation, especially when the final assemblages were dominated by diatoms. Synechococcus sp. were not grazed and only showed moderate to no response to nutrient additions. The high spatial and temporal variation of our results indicates that the composition of the planktonic community determines the prevailing control mechanisms. It further implies that, at this transitional time of the year (onset of summer stratification), the populations fluctuate about an equilibrium between growth and grazing.Communicated by O. Kinne, Oldendorf/Luhe     

Seasonal variations in the growth rates of euphausiids (<Emphasis Type="Italic">Thysanoessa inermis</Emphasis>, <Emphasis Type="Italic">T. spinifera</Emphasis>, and <Emphasis Type="Italic">Euphausia pacifica</Emphasis>) from the northern Gulf of Alaska

The euphausiids Thysanoessa inermis (Kroyer 1846), Thysanoessa spinifera (Holmes 1900), and Euphausia pacifica (Hansen 1911) are key pelagic grazers and also important prey for many commercial fish species in the Gulf of Alaska (GOA). To understand the role of the euphausiids in material flows in this ecosystem their growth rates were examined using the instantaneous growth rate (IGR) technique on the northern GOA shelf from March through October in 2001–2004. The highest mean molting increments (over 5% of uropod length increase per molt) were observed during the phytoplankton bloom on the inner shelf in late spring for coastal T. inermis, and on the outer shelf in summer for T. spinifera and more oceanic E. pacifica, suggesting tight coupling with food availability. The molting rates were higher in summer and lower in spring, for all species and were strongly influenced by temperature. Mean inter-molt periods calculated from the molting rates, ranged from 11 days at 5°C to 6 days at 8°C, and were in agreement with those measured directly during long-term laboratory incubations. Growth rate estimates depended on euphausiid size, and were close to 0 in early spring, reaching maximum values in May (0.123 mm day⁻¹ or 0.023 day⁻¹ for T. inermis) and July (0.091 mm day⁻¹ or 0.031 day⁻¹ for T. spinifera). The growth rates for E. pacifica remained below 0.07 mm day⁻¹ (0.016 day⁻¹) throughout the season. The relationship between T. inermis weight specific growth rate (adjusted to 5°C) and ambient chlorophyll-a concentration fit a Michaelis–Menten curve (r ² = 0.48) with food saturated growth rate of 0.032 day⁻¹ with half saturation occurring at 1.65 mg chl-a m⁻³, but such relationships were not significant for T. spinifera or E. pacifica.     

Growth and development of nauplii and copepodites of the estuarine copepod Acartia tonsa from southern Europe (Ria de Aveiro, Portugal) under saturating food conditions

A temperature-dependent growth model is presented for nauplii and copepodites of the estuarine calanoid copepod Acartia tonsa from southern Europe (Portugal). Development was followed from egg to adult in the laboratory at four temperatures (10, 15, 18 and 22°C) and under saturating food conditions (>1,000 μg C l⁻¹). Development times versus incubation temperature were fitted to a Belehradek’s function, showing that development times decreased with increasing incubation temperature: at 10°C, A. tonsa need 40.3 days to reach adult stage, decreasing to 8.9 days when reared at 22°C. ANCOVA (homogeneity of slopes) showed that temperature (P<0.001) and growth phase (P<0.01) had a significant effect on the growth rate. Over the range of temperatures tested in this study, highest weight-specific growth rates were found during naupliar development (NI–NVI) and varied from 0.185 day⁻¹ (10°C) to 0.880 day⁻¹ (22°C) with a Q ₁₀ equal to 3.66. During copepodite growth (CI–CV), the weight-specific growth rates ranged from 0.125 day⁻¹ (10°C) to 0.488 day⁻¹ (22°C) with a Q ₁₀ equal to 3.12. The weight-specific growth rates (g) followed temperature (T) by a linear relationship and described as ln g=−2.962+0.130 T (r ²=0.99, P<0.001) for naupliar stages and ln g=−3.134+0.114T (r ²=0.97, P<0.001) for copepodite stages. By comparing in situ growth rates (juvenile growth and fecundity) for A. tonsa taken from the literature with the temperature-dependent growth model defined here we suggest that the adult females of A. tonsa are more frequently food limited than juveniles.