Spring bloom sedimentation in a subarctic ecosystem

A 5-yr study (1985 to 1989) of spring bloom sedimentation in Auke Bay, Alaska, indicates that the sinking response of diatoms to ambient nutrients influences both species succession during the spring bloom and the subsequent sedimentation of new production. Diatoms from the genera Thalassiosira, Chaetoceros and Skeletonema formed the bulk of the spring bloom each year. Growth of Thalassiosira spp. consistently initiated the primary bloom, while Skeletonema costatum tended to grow later in, or after, the primary bloom. We postulate that this successional pattern is driven by interspecific nutrient competition. Overall, sedimentation flux of the dominant species of bloom diatoms was correlated with surface concentrations of cells integrated over the bloom period. In fact, different linear relationships existed when Thalassiosira and Chaetoceros spp. were considered separately, but not for Skeletonema sp., indicating that marked differences exist between the sedimentation tendencies of these genera. The observed inter-generic differences are explicable by the different overall sinking rates, as well as different nutrient-sensitivities of the sinking rates of each genus. Thalassiosira spp., the fastestsinking and most nutrient-sensitive species, contributed up to 10 x more carbon to the benthos in all years of the study, reaching a maximum of 11.1 gCm^-2 over a single spring bloom event in 1988. This study indicates that the tendency to sink to the benthos during and/or after a bloom is highly dependent on species-specific cell physiology, and supports the idea that it is the fast-sinking, nutrient-sensitive diatoms, such as Thalassiosira species, that constitute the major source of vertical carbon flux in this embayment and other such coastal ecosystems during the spring bloom.     

Species-specific sedimentation and sinking velocities of diatoms

Sedimentation rates were determined for various diatom species, and both average and maximum sinking velocities of sedimenting diatoms were calculated during a spring bloom investigation in the central Baltic Sea in 1986. Up to 25 and 50% of theChaetoceros spp. andThalassiosira levanderi populations, respectively, sedimented daily. Daily sedimentation rates of other diatoms, dinoflagellates andMesodinium rubrum, however, were less than 1% of their respective standing stocks. TheT. levanderi population was divided into two subpopulations: while one was sinking, the second was actively dividing (recognizable by paired-cell stages) with a specific growth rate of >0.2 to 0.3 d^–1. These paired cells were never found in sediment trap samples. The average sinking velocity ofChaetoceros spp. was 15 to 30 m d^–1; that ofT. levanderi was higher. The maximum sinking velocity of cells was at least 70 m d^–1. According to these observations, the formation of aggregates (which enhances sinking velocity), and their sedimentation, represent a highly selective process. This indicates that diatom aggregates do not act as roving filters, sweeping the water clear while sinking.     

High reproduction of <Emphasis Type="Italic">Calanus finmarchicus</Emphasis> during a diatom-dominated spring bloom

Feeding, egg production, hatching success and early naupliar development of Calanus finmarchicus were measured in three north Norwegian fjords during a spring bloom dominated by diatoms and the haptophyte Phaeocystis pouchetii. Majority of the copepod diet consisted of diatoms, mainly Thalassiosira spp. and Chaetoceros spp., with clearance rates up to 10 ml ind⁻¹ h⁻¹ for individual algae species/groups. Egg production rates were high, ranging from ca 40 up to 90 eggs f⁻¹ d⁻¹, with a hatching success of 70–85%, and fast naupliar development through the first non-feeding stages. There was no correlation between the egg or nauplii production and diatom abundance, but the hatching success was slightly negatively correlated with diatom biomass. However, the overall high reproductive rates suggested that the main food items were not harmful for C. finmarchicus reproduction in the area, although direct chemical measurements were not conducted. The high population egg production (>1,20,000 eggs m⁻² d⁻¹) indicated that a large part of the annual reproduction took place during the investigation, which stresses the importance of diatom-dominated spring phytoplankton bloom for population recruitment of C. finmarchicus in these northern ecosystems.     

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%).     

Coagulation efficiency and aggregate formation in marine phytoplankton

Flocculation of phytoplankters into large, rapidly sinking aggregates has been implicated as a mechanism of vertical transport of phytoplankton to the sea floor which could have global significance. The formation rate of phytoplankton aggregates depends on the rate at which single cells collide, which is mainly physically controlled, and on the probability of adhesion upon collision (=coagulation efficiency, stickiness), which depends on physico-chemical and biological properties of the cells. We describe here an experimental method to quantify the stickiness of phytoplankton cells and demonstrate that three species of diatoms grown in the laboratory (Phaeodactylum tricornutum, Thalassiosira pseudonana, Skeletonema costatum) are indeed significantly sticky and form aggregates upon collision. The dependency of stickiness on nutrient limitation and growth was studied in the two latter species by investigating variation in stickiness as batch cultures aged. In nutrient repleteT. pseudonana cells stickiness is very low (< 5 × 10^?3), but increases by more than two orders of magnitude as cell growth ceases and the cells become nutrient limited. Stickiness ofS. costatum cells is much less variable, and even nutrient replete cells are significantly sticky. Stickiness is highest (> 10^?1) forS. costatum cells in the transition between the exponential and the stationary growth phase. The implications for phytoplankton aggregate formation and subsequent sedimentation in the sea of these two different types of stickiness patterns are discussed.     

Effects of light quality on initiation and development of meroplanktonic diatom blooms in a eutrophic shallow sea

We investigated the effects of light quality on resting stage cell germination and vegetative cell growth of meroplanktonic diatoms in a small port in Hakata Bay, Japan and in the laboratory. During the investigation over the year of 2006, the meroplanktonic diatom bloom first occurred in the end of May and then repeated wane and wax until October in the small port. From late April to middle May, light penetrating the water column was often strong and attenuations of all spectral lights were low. During this period, Skeletonema costatum, Thalassiosira minima, and Chaetoceros sp. appeared frequently, followed by the blooms of S. costatum and Chaetoceros sp. in late May. Thereafter, S. costatum and Chaetoceros sp. bloomed in late June but not in middle June, when pigmented flagellates bloom appeared. The attenuation of short-wavelength light such as violet and blue lights was markedly high during these diatom and flagellate blooms; all blooms disappeared within several days. Vegetative cell strains of the three diatoms under light emitting diodes (LEDs) with six different spectra (violet, blue, green, orange, red, and near-infrared) grew at a higher rate under short-wavelength light, violet and blue. On the other hand, when suspensions of bottom sediments from Hakata Bay were cultured under the same LEDs and in the dark, vegetative cells of S. costatum appeared under all LEDs except for orange and near-infrared, vegetative cells of T. minima appeared under all LEDs but not in the dark, and vegetative cells of Chaetoceros sp. appeared under violet and blue LEDs. However, vegetative cell densities of the three diatoms increased much more under violet light than under other LEDs within a short period (6 days). Our study indicates that underwater penetration by short-wavelength light, such as violet and blue, may be an important factor in the initiation and development of meroplanktonic diatom blooms.     

The role of ciliates,heterotrophic dinoflagellates and copepods in structuring spring plankton communities at Helgoland Roads,North Sea

Mesocosm experiments coupled with dilution grazing experiments were carried out during the phytoplankton spring bloom 2009. The interactions between phytoplankton, microzooplankton and copepods were investigated using natural plankton communities obtained from Helgoland Roads (54°11.3′N; 7°54.0′E), North Sea. In the absence of mesozooplankton grazers, the microzooplankton rapidly responded to different prey availabilities; this was most pronounced for ciliates such as strombidiids and strobilids. The occurrence of ciliates was strongly dependent on specific prey and abrupt losses in their relative importance with the disappearance of their prey were observed. Thecate and athecate dinoflagellates had a broader food spectrum and slower reaction times compared with ciliates. In general, high microzooplankton potential grazing impacts with an average consumption of 120% of the phytoplankton production (P _p ) were measured. Thus, the decline in phytoplankton biomass could be mainly attributed to an intense grazing by microzooplankton. Copepods were less important phytoplankton grazers consuming on average only 47% of P _p . Microzooplankton in turn contributed a substantial part to the copepods’ diets especially with decreasing quality of phytoplankton food due to nutrient limitation over the course of the bloom. Copepod grazing rates exceeded microzooplankton growth, suggesting their strong top-down control potential on microzooplankton in the field. Selective grazing by microzooplankton was an important factor for stabilising a bloom of less-preferred diatom species in our mesocosms with specific species (Thalassiosira spp., Rhizosolenia spp. and Chaetoceros spp.) dominating the bloom. This study demonstrates the importance of microzooplankton grazers for structuring and controlling phytoplankton spring blooms in temperate waters and the important role of copepods as top-down regulators of microzooplankton.     

Evaluation of fatty acids as biomarkers for a natural plankton community. A field study of a spring bloom and a post-bloom period off West Greenland

An investigation of the fatty acid composition of a natural arctic plankton community was carried out over two fishing banks located between 63°N and 65°N off the West Greenland coast. Samples for fatty acid analyses, species determination and biomass assessments of the plankton community were taken at the depth of fluorescence maximum. High biomass and diatom dominance during the spring bloom and low biomass and flagellate dominance in the post-bloom period were reflected by the fatty acid profiles. The total amount of fatty acid ranged from 55 to 132 µg l^-1 during the spring bloom and from 1 to 5 µg l^-1 during the post bloom. Analysis of the fatty acids showed that when the plankton was dominated by diatoms of the genera Thalassiosira and Chaetoceros, the proportions of C16:1(n-7) and C20:5(n-3) were correspondingly high. C18s, and particularly C18:1(n-9), were more abundant when the plankton was dominated by small autotrophic flagellates, primarily haptophytes. We found a good positive correlation between the common diatom marker, C16:1(n-7)/C16:0, and the biomass percentage of diatoms (r=0.742, P<0.001), as well as between the biomass percentage of flagellates and total C18 fatty acids (r=0.739, P<0.001). This supports the use of these specific fatty acids and fatty acid ratios as general biomarkers of the plankton community. However, the fatty acids are not specific enough to sufficiently characterise the composition of the plankton community, and microscopical support is needed to verify observed trends.     

Sinking rate response to depletion of nitrate,phosphate and silicate in four marine diatoms

The effect of nutrient (N, P, and Si) depletion on sinking rates was studied for two small (Skeletonema costatum (Grev.) Cleve and Chaetoceros gracile Schütt) and two large [Ditylum brightwellii (West) Grun and Coscinodiscus wailesii (Gran et Angst)] centric diatoms obtained from stock cultures. Each diatom was examined under conditions of (1) nutrient repletion (=log growth phase), (2) nutrient depletion (48 h without a given substrate), and (3) recovery (24h after addition of limiting substrate to nutrient-deplete populations). All nutrient-replete cultures displayed low sinking rates despite large differences in cell size. In nutrient-deplete populations, sinking rate was related to the kind of nutrient depleted and varied among species. Silicate depletion elicited by far the greatest increase in sinking rates in all 4 species, indicating that biochemical aspects of silicon metabolism are more important to buoyancy regulation than density-related variations in the amount of silicon per cell. Since N- and P-depletion caused lower sinking rates in 3 of the species, this observation calls for re-evaluation of the axiom that nutrient depletion necessarily causes increased sinking rates. The exception was Coscinodiscus wailesii, which sank faster under all types of nutrient limitation. In most cases, sinking rates typical of log-phase cultures were not regained within 24 h after the addition of limiting nutrient to nutrient-depleted populations. Ultimately, the length of the recovery period may be useful in identifying the metabolic processes responsible for buoyancy regulation in actively growing cells.     

Primary production and phytoplankton community structure during a winter shelf-scale phytoplankton bloom off Western Australia

Regions of high primary production along the oligotrophic west coast of Australia between 34 and 22°S in May–June 2007 (midway through the annual phytoplankton bloom) were found around mesoscale features of the Leeuwin Current. At 31°S, an anticyclonic eddy-forming meander of the Leeuwin Current had a mixed layer depth of >160 m, a depth-integrated chlorophyll a (Chl a)-normalised primary production of 24 mg C mg Chl a ^?1 day^?1 compared to the surrounding values of <18 mg C mg Chl a ^?1 day^?1. In the north between 27 and 24°S, there were several stations in >1,000 m of water with a shallow (<100 m) and relatively thin layer of high nitrate below the mixed layer but within the euphotic zone. These stations had high primary production at depths of ~100 m (up to 7.5 mg C m^?3 day^?1) with very high rates of production per unit Chl a (up to 150 mg C mg Chl a ^?1 day^?1). At 27–24°S, the majority of the phytoplankton community was the ubiquitous tropical picoplankters, Synechococcus and Prochlorococcus. There was a decline in the dominance of the picoplankters and a shift towards a more diverse community with more diatoms, chlorophytes, prasinophytes and cryptophytes at stations with elevated production. Photosynthetic dinoflagellates were negligible, but heterotrophic dinoflagellate taxa were common. Haptophytes and pelagophytes were also common, but seemed to contribute little to the geographical variation in primary production. The mesoscale features in the Leeuwin Current may have enhanced horizontal exchange and vertical mixing, which introduced nitrate into the euphotic zone, increasing primary production and causing a shift in phytoplankton community composition in association with the annual winter bloom.     

Sinking-rate response of natural assemblages of temperate and subtropical phytoplankton to nutrient depletion

Phytoplankton assemblages were collected during spring blooms in 1982 in Washington State and in Hawaii. Sinking rate responses of these assemblages were examined under nitrate, phosphate, and silicate depletion. Ambient nutrient concentrations, chlorophyll concentrations, photosynthetic rates, sinking rates, and floristic compositions were determined. Under nutrient-replete conditions, the temperate assemblage, composed primarily of large centric diatoms, had a sinking rate of 0.96 m d^-1; sinking rates did not change appreciably over 4 d without nitrate. Without phosphate or silicate, the sinking rates remained constant for 3 d and then increased after biomass indices began to decline. These findings illustrate the potential importance of phosphate or silicate depletion to the sedimentation of spring-bloom diatom populations. The subtropical assemblage, composed primarily of diatoms, coccolithophorids, and dinoflagellates, had an initial sinking rate of 0.22 m d^-1 and did not display substantial sinking rate changes in the absence of nitrate, phosphate or silicate. Floristic data consistently showed a proliferation of pennate diatoms, which had lower settling rates than centric diatoms. Growth and sedimentation patterns indicated a competitive advantage for pennate diatom components of subtropical assemblages; this in turn may limit phytoplankton sedimentation losses in such ecosystems.     

Seasonal studies on the relative importance of different size fractions of phytoplankton in Narragansett Bay (USA)

The composition and productivity of four different size-fractions (<20, 20 to 60, 60 to 100, >100 μm) of the phytoplankton of lower Narragansett Bay (USA) were followed over an annual cycle from November, 1972 to October, 1973. Diatoms dominated the population in the winter-spring bloom and in the fall, the summer population was dominated by flagellates. The nannoplankton (<20 μm) were the most important, accounting for 46.6% of the annual biomass as chlorophyll a and 50.8% of the total production. The relative importance of the different fractions showed a marked seasonality. During the winter-spring and fall blooms the netplankton fractions (>20 μm) were the most important. Nannoplankters domnated in the summer. The yearly mean assimilation numbers for the different fractions were not signfficantly different. During the winter-spring bloom, however, the assimilation numbers for the netplankters were significantly higher than those for the nannoplankton fraction. Temperature accounted for most of the variability in assimilation numbers; a marked nutrient stress was observed on only two occasions. Growth rates calculated from ¹⁴C uptake and adenosine triphosphate (ATP)-cell carbon were generally quite high; maxima were >1.90 doublings per day during blooms of a flagellate in the summer and of Skeletonema costatum in the fall. The series of short cycles observed in which the dominant species changed were related to changes in the physiological state of the population. Higher growth rates were generally observed at times of peak phytoplankton abundance while lower growth rates were observed between these peaks. The high growth rates and assimilation numbers usually found suggest that the phytoplankton in lower Narragansett Bay was not generally nutrient-limited between November, 1972 and October, 1973. Nutrient regeneration in this shallow estuary, therefore, must be very rapid when in situ nutrient levels are low.     

Buoyancy of natural populations of marine phytoplankton

Buoyancy of natural populations of marine phytoplankton was studied in a fjord in western Norway during the diatom bloom and in autumn. The study was carried out under approximate in situ conditions by means of an apparatus described in the paper. During the spring bloom, positive buoyancy was observed only once. Sinking rates of individual fractions ranged from 0 to more than 9 m day^-1, and the mean sinking rates of the total chlorophyll content from 0 to at least 2. 2 m day^-1. The highest rates occurred in the post-bloom period, while sinking appeared negligible from the onset of the bloom up to its culmination. In autumn, the population was dominated by small, flagellated cells. Positive buoyancy, or upward migration, was then observed in two out of three experiments.     

Quantifying plant pigments and live diatoms in aphotic sediments of Scandinavian coastal waters confirms a major route in the pelagic–benthic coupling

The fate of pelagic diatoms in marine coastal aphotic sediments was investigated from sediment profiles in western Scandinavian waters. We used three independent methods to estimate pigment pools in the sediment: (1) fluorometry, (2) high-performance liquid chromatography and (3) pigments estimated from germinable diatom cells, using the dilution extinction method. A strong positive relationship with an intercept close to zero was observed between fucoxanthin, a marker of diatoms, and chlorophyll a. The fucoxanthin/chl a ratio was on average 1.05, which was similar to monocultures of dominating diatom taxa, indicating that sedimentary chl a was to a large extent of diatom origin. Chl a and fucoxanthin correlated significantly and positively with, and where within the same order of magnitude as, corresponding substances predicted from live diatom cell numbers obtained with the dilution extinction method. This indicates that a major part of surficial sediment chl a was bound in live cells of pelagic diatoms. There was a consistent change in viable cells with sediment depth and with timing of dominating taxa, with the non-spore-forming Skeletonema costatum dominating in the surface sediment in March and May, while the spore forming Chaetoceros spp. dominated deep in the sediment and during periods outside of the spring bloom (February and August). This indicates that chl a is bound in several different cell pools with different degradation rates, depending on diatom taxonomy. Thus, diatoms originating predominantly from the spring bloom may provide an important direct link in the pelagic–benthic coupling in this area.     

Comparison between chemical and isotopic measurements of biological nitrate utilization: further evidence of low new-production levels in the equatorial Pacific

New-production (nitrate uptake) rates in the equatorial Pacific were estimated by parallel measurements of nitrate disappearance from sea water using a colorimetric method and of ¹⁵N-labelled nitrate (¹⁵NO₃ ⁻) incorporation into particulate organic nitrogen (PON) collected on GF/F filters (net nitrate uptake, conventional ¹⁵N-tracer method) and Anopore (0.2 μm) membranes. Regression analyses of 74 sample pairs gathered during 12 and 24 h productivity experiments revealed a significant positive relationship between decreasing nitrate level and ¹⁵NO₃ ⁻ accumulation into PON retained on GF/F filters, but the slopes of Model I and Model II regression lines were 1.18 and 1.29, respectively, suggesting that 15 to 22% of ¹⁵NO₃ ⁻ removed from the dissolved fraction were lost to another N-pool. Two possible avenues for the missing ¹⁵NO₃ ⁻ have been examined: uptake by submicron particles passed through the GF/F filters, and loss as dissolved organic nitrogen (DON). Nitrate uptake by small cells not recovered on GF/F filters, could be safely eliminated as a cause of loss, since ¹⁵NO₃ ⁻ uptake rates obtained from ¹⁵N entering PON collected on GF/F filters agreed well with those obtained from ¹⁵N entering PON collected on Anopore membranes (32 sample pairs). Inspection of the DON pool of 0.2 μm filtrates for excess-¹⁵N enrichment (20 samples) revealed that in nitrate-rich waters (equatorial upwelling between 1°N and 10°S), loss of ¹⁵NO₃ ⁻ as DO¹⁵N accounted for <5% of net nitrate uptake. In samples from subtropical oligotrophic waters (from 11°S southward), however, ¹⁵NO₃ loss as DO¹⁵N represented up to 20% of net NO₃ ⁻ uptake. These results, as well as experimental considerations concerning the use of colorimetric and isotopic methods to measure new production show that: (1) earlier reported high discrepancies between nitrate decreases (ΔNO₃ ⁻) and ¹⁵NO₃ ⁻ incorporation into filterable particles (ΔNO₃ ⁻/¹⁵NO₃ ⁻ incorporation >2) were probably erroneous; (2) the use of GF/F filters does not result in an underestimation of new production, although it was found to underestimate PON concentrations by up to 60%; (3) in the equatorial upwelling area (1°N to 10°S), which has high ambient nitrate levels (>2000 nmol l⁻¹) but only slight changes in concentration (0 to 80 nmol l⁻¹ d⁻¹), new production is more accurately estimated by the isotopic method than by the chemical method; (4) in subtropical oligotrophic waters (from 11°S southward) with low ambient nitrate levels (0 to 100 nmol l⁻¹), both procedures are appropriate as long as nitrate removal per incubation period is >3 nmol l⁻¹ (lower rates are only detectable with the isotopic method); (5) the traditional ¹⁵N-tracer technique does not substantially underestimate net new-production in the equatorial Pacific, and failure to account for the loss of ¹⁵NO₃ ⁻ as DON, i.e. to estimate gross nitrate uptake (gross uptake = net uptake + ¹⁵N loss) tends to underestimate new production on an average by only 10%. Overall, the apparent low level of new production in the nitrate-rich area of the central equatorial Pacific seems to be a fact, and may be ascribable to other nutrient (macro and micro) deficiencies and/or to intense in situ recycling of ammonium and nitrate (regenerated production) rather than to inaccurate nitrate uptake rates measured with the classical ¹⁵N-tracer technique. Received: 24 November 1998 / Accepted 10 March 2000     

“Seed bank” of coastal planktonic diatoms in bottom sediments of Hiroshima Bay, Seto Inland Sea, Japan

Abundance and temporal distribution of viable (able to germinate) resting stage cells of planktonic diatoms in bottom sediments have been investigated almost monthly during 1989 to 1992 in Hiroshima Bay, western part of Seto Inland Sea, Japan. The abundance of viable resting stages in bottom sediments was enumerated with the extinction dilution method (most probable number method, MPN). In bottom sediments of Hiroshima Bay, dominantly distributed species and/ or genera of the diatom resting stages were Skeletonema costatum, Chaetoceros spp. and Thalassiosira spp. Viable resting stages of these diatoms were densely distributed on the orders of 10³ to 10⁶ (MPN g⁻¹ wet sediments), and persisted in bottom sediments throughout the investigation period. Conversely, vegetative cells of these diatoms fluctuated remarkably in the water column and disappeared sporadically. Survival of the resting stages in a collected sediment sample was also determined with the MPN method, at different storage temperatures (5, 10, 15, 20, 25 °C). The survival test demonstrated that the diatom resting stages could survive in the dark for several months or years in sediments. Resting stages survived longer at the lower storage temperature, and the order of longevity was consistent within three diatoms (Chaetoceros spp. > Thalassiosira spp. > S. costatum) at each storage temperature. The present study suggests that these diatom resting stages in the coastal bottom sediments could serve as a “seed bank”, analogous to those of terrestrial plants. The seed bank would ensure the survival of diatoms within highly fluctuating coastal environments, while it would also be the source of sporadic and autochthonous diatom blooms in coastal waters. Received: 29 November 1996 / Accepted: 16 December 1996     

Interactions between nitrate and ammonium uptake: variation with growth rate,nitrogen source and species

The interaction between nitrate and ammonium uptake was examined as a function of preconditioning growth rate and nitrogen source by adding nitrate, ammonium, or both to nitrogen-sufficient,-deficient, and-starvedSkeletonema costatum (Grev.) Cleve and nitrogen-deficientChaetoceros debilis Cleve. By simultaneously measuring the internal accumulation of intermediates of nitrogen assimilation and the rates of nitrogen assimilation, the metabolic control of nitrogen uptake could be assessed. After the simultaneous addition of nitrate and ammonium to culture, both nitrate and ammonium uptake rates were decreased in comparison with the rates observed when each was added alone, although nitrate uptake was usually decreased more than ammonium uptake. Since both nitrate and ammonium uptake rates vary with time, preconditioning growth conditions, nitrogen sources present, and species, it was necessary to use several different indices to quantify inhibition. In general, ammonium inhibition of nitrate uptake inS. costatum was greatest in cultures preconditioned to ammonium and those at low growth rates, whereas ammonium uptake was inhibited most in cultures preconditioned to nitrate. In nitrogen-deficientC. debilis, nitrate uptake was more inhibited by ammonium, but uptake returned to normal rates more quickly than inS. costatum, whereas inhibition of ammonium uptake was similar. These results explain why the interaction between nitrate and ammonium uptake in the field can be so variable. Inhibition of uptake is not controlled by internal ammonium or total amino acids, nor is it related to the inability to reduce nitrate. Instead, inhibition must be determined in part by the external concentration of nitrogen compounds and in part by some intermediate(s) of nitrogen assimilation present inside the cell.Bigelow Laboratory Contribution No. 82022     

Long-term changes in phytoplankton phenology and community structure in the Bahía Blanca Estuary, Argentina

The phytoplankton of the Bahía Blanca Estuary, Argentina, has been surveyed since 1978. Chlorophyll a, phytoplankton abundance, species composition and physico-chemical variables have been fortnightly recorded. From 1978 to 2002, a single winter–early spring diatom bloom has dominated the main pattern of phytoplankton interannual variability. Such pattern showed noticeable changes since 2006: the absence of the typical winter bloom and changes in phenology, together with the replacement of the dominant blooming species, i.e. Thalassiosira curviseriata, and the appearance of different blooming species, i.e. Cyclotella sp. and Thalassiosira minima. The new pattern showed relatively short-lived diatom blooms that spread throughout the year. In addition, shifts in the phytoplankton size structure toward small-sized diatoms, including the replacement of relatively large Thalassiosira spp. by small Cyclotella species and Chaetoceros species have been noticed. The changes in the phenology and composition of the phytoplankton are mainly attributed to warmer winters and the extremely dry weather conditions evidenced in recent years in the Bahía Blanca area. Changing climate has modified the hydrological features in the inner part of the estuary (i.e. higher temperatures and salinities) and potentially triggered the reorganization of the phytoplankton community. This long-term study provides evidence on species-specific and structural changes at the bottom of the pelagic food web likely related to the recent hydroclimatic conditions in a temperature estuary of the southwestern Atlantic.     

The rapid response of the marine diatom Skeletonema costatum to changes in external and internal nutrient concentration

A nitrogen-deficient batch culture of the marine diatom Skeletonema costatum, when resupplied with a mixture of nitrate and ammonium, showed an initial enhanced nitrate uptake rate leading to a large internal concentration (pool) of nitrate. Following this initial nitrate uptake event, nitrate uptake ceased, and nitrate assimilation was inhibited until the ammonium present was used. At this point, nitrate uptake resumed and nitrate assimilation began. No internal ammonium pool was observed during nitrate utilization, but a large nitrate pool remained throughout the utilization of external nitrate. The internal nitrate pool decreased rapidly after exhaustion of nitrate from the culture medium, but growth of cellular particulate nitrogen continued for about 24 h. A mathematical simulation model was developed from these data. The model cell consisted of a nitrate pool, ammonium pool, dissolved organic nitrogen pool, and particulate nitrogen. It was found that simple Michaelis-Menten functions for uptake and assimilation gave inadequate fit to the data. Michaelis-Menten functions were modified by inclusion of inhibitory and stimulatory feedback from the internal pools to more accurately represent the observed nutrient utilization.     

Contribution of salps to carbon flux of marginal ice zone of the Lazarev Sea, southern ocean

In order to estimate the in situ grazing rates of Salpa thompsoni and their implications for the development of phytoplankton blooms and for the sequestration of biogenic carbon in the high Antarctic, a repeat-grid survey and drogue study were carried out in the Lazarev Sea during austral summer of 1994/1995 (December/January). Exceptionally high grazing rates were measured for S. thompsoni at the onset of a phytoplankton bloom (0.2 to 0.8 μg chlorophyll a l⁻¹) in December 1994, with up to ≃160 μg of plant pigments consumed by an individual salp of 7 to 10 cm length per day. Dense salp swarms extended throughout the marginal ice zone, consuming up to 108% of daily phytoplankton production and 21% of the total chlorophyll a stock. Due to the much faster sinking rates and higher carbon content of salp faecal pellets, the efficiency of downward carbon flux through salps is much higher than through the other major grazers, krill and copepods. S. thompsoni can thus export large amounts of biogenic carbon from the euphotic zone to the deep ocean. With the observed ingestion rates during December 1994, this flux could have attained levels of up to 88 mg C m⁻² d⁻¹, accounting for the bulk of the vertical transport of carbon in the Lazarev Sea. However, in January 1995, when phytoplankton concentrations exceeded a threshold level of 1.0 to 1.5 μg chlorophyll a l⁻¹, salps experienced a drastic reduction in their feeding efficiency, possibly as a result of clogging of their filtering apparatus. This triggered a dramatic reversal in the relationship, during which a dense phytoplankton bloom developed in conjunction with the collapse of the salp population. Increases in the biomass and geographic range of the tunicate S. thompsoni have occurred in several areas of the southern ocean, often in parallel with a rise in sea-surface temperature during sub-decadal periods of warming anomalies. Received: 10 August 1997 / Accepted: 21 October 1997