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.     

Role of sinking in diatom life-history cycles: ecological,evolutionary and geological significance

Rapid mass sinking of cells following diatom blooms, observed in lakes and the sea, is argued here to represent the transition from a growing to a resting stage in the life histories of these algae. Mass sinking is of survival value in those bloom diatoms that retain viability over long periods in cold, dark water but not in warm, nutrient-depleted surface water. Mechanisms for accelerating sinking speed of populations entering a resting or seeding mode are proposed. Previously unexplained features of diatom form and behaviour take on a new meaning in this context of diatom seeding strategies. Diatoms have physiological control over buoyancy as declining growth is accompanied by increasing sinking rates, where the frustule acts as ballast. Increased mucous secretion in conjunction with the cell protuberances characteristic of bloom diatoms leads to entanglement and aggregate formation during sinking; the sticky aggregates scavenge mineral and other particles during descent which further accelerates the sinking rate. Such diatom flocs will have sinking rates of 100 m d^-1 or more. This is corroborated by recent observations of mass phytoplankton sedimentation to the deep sea. This mechanism would explain the origin of marine snow flocs containing diatoms in high productivity areas and also the well-known presence of a viable deep sea flora. That mortality is high in such a seeding strategy is not surprising. A number of species-specific variables pertaining to size, morphology, physiology, spore formation and frustule dissolution rate will determine the sinking behaviour and thus control positioning of resting stages in the water column or on the bottom. It is argued that sinking behaviour patterns will be environmentally selected and that some baffling aspects of diatom form and distribution can be explained in this light. Rapid diatom sedimentation is currently believed to be mediated by zooplankton faecal pellets, particularly those of copepods. This view is not supported by recently published observations. I speculate that copepod grazing actually retards rather than accelerates vertical flux, because faecal pellets tend to be recycled within the surface layer by the common herbivorous copepods. Egestion of undigested food by copepods during blooms acts as a storage mechanism, as ungrazed cells are likely to initiate mass precipitation and depletion of the surface layer in essential elements. Unique features of diatoms are discussed in the light of their possible evolution from resting spores of other algae. An evolutionary ecology of pelagic bloom diatoms is deduced from behavioural and morphological characteristics of meroplanktonic and tychopelagic forms. Other shell-bearing protistan plankters share common features with diatoms. Similar life-history patterns are likely to be present in species from all these groups. The geological significance of mass diatom sinking in rapidly affecting transfer of biogenic and mineral particles to the sea floor is pointed out.     

Size structure of phytoplankton biomass and photosynthesis in subtropical Hawaiian waters

In a subtropical Hawaiian ecosystem, phytoplanton size structure analyses (November–December, 1980) showed that ultraplankton (>3μm), nanoplankton (>20μm) and netplankton (>20μm) accounted for ca. 80, 98, and 2% of total chlorophyll standing stock, respectively, on the basis of chlorophyll. Similar trends were evident, for other biomass indices (e.g. cell numbers, total cell volume, ATP, particulate organic carbon, particulate organic nitrogen). The ultraplankton fraction consisted primarily of small flagellates (1 to 3 μm diam) and coccoid cells (?1 μm diam); the 3 to 20 μm fraction was represented by dinoflagellates, coccolithophores, diatoms, and chrysophytes; and the netplankton fraction consisted principally of dinoflagellates and centric diatoms. Community photosynthesis had a size distribution similar to that of biomass. Sinking rates for the 3 μm, 3 to 20 μm, and >20 μm fractions averaged 0.0, 0.09, and 0.29m d^?1, respectively. The absence of measurable sinking rates for the ultraplankton, together with the relative abundance of biomass in this fraction, result in very small phytoplankton losses due to sinking in such subtropical surface waters.     

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.     

Growth rates of summer nanoplankton (<10 μm) populations in lower Narragansett Bay,Rhode Island,USA

Growth rates of summer (June–September) phytoplankton assemblages and constituent species were measured in 30 diffusion culture experiments. Size-fractionated (<10 m) phytoplankton assemblages were incubated in situ or under simulated in-situ conditions in outdoor tanks connected to a running seawater system. Doubling rates of important species and groups (such as microflagellates) were compared to community biomass doubling rates estimated from ¹⁴C uptake and changes in chlorophyll a concentrations. Division rates of dominant diatom species generally equalled or exceeded community biomass doubling rates, while those of flagellates and non-motile ultraplankters were slower. Maximum division rates of sixteen common diatom species exceeded 2.1 divisions d^-1, while nine had maximum division rates in excess of 3 d^-1. Mean division rates of 12 diatom species exceeded 1 d^-1. Maximum division rates of flagellated species, uncharacterized microflagellates and non-motile ultraplankton assemblages were 2.1, 1.5 and 1.4 d^-1, respectively. Microflagellate and non-motile ultraplankton assemblage doubling rates were less than 0.5 d^-1 in over half of all growth experiments.     

Phytoplankton in the Weddell Sea,Antarctica: composition,abundance and distribution in water-column assemblages of the marginal ice-edge zone during austral autumn

At the compacted, north-south line of the ice edge, phytoplankton were sampled during early austral autumn of 1986 in the northwestern Weddell Sea. Cells from discrete water bottle samples from 12 stations on two east-west transects were counted to gain quantitative information on the composition, abundance, distribution, and condition of the phytoplankton in water-column assemblages. Over 70 species were found. The highest numbers of total cells (integrated through the top 150 m) were found in open water, well-separated from and to the east of the ice edge on the southern transect, with 6.01×10¹⁰ cells m^-2. The relative abundance of diatoms was low at ice-convered stations (< 35% of the total phytoplankton in preserved samples) and high at open-water stations (> 80%); however, the relative abundance of the prymnesiophyte Phaeocystis sp. was high at ice-covered stations (> 60%) and low at open-water stations (< 16%), with lower absolute abundances than during a previous austral-spring phytoplankton increase. In the open ocean, the dominants were the pennate diatoms Fragilariopsis cylindrus, Pseudonitzschia prolongatoides, F. curta, and a small form of the centric diatom Chaetoceros dichaeta in chains. Although the three pennate diatoms were frequently dominant in number, they represented less biomass than C. dichaeta in open waters. Mean phytoplankton abundance was low (0.2×10⁶ cells l^-1) but, overall, the diatom cell density (0.14×10⁶ cells l^-1) was similar to that found previously during a northward transect from ice-covered to ice-free water at the Weddell-Scotia Sea ice edge (spring 1983). The phytoplankton spatial patterns in the two autumn transects differed, with the more southerly transect exhibiting a higher abundance of diatoms and dinoflagellates. The ratio of full to empty diatoms was higher on the southern transect, indicating a healthy population, while lower ratios of full/empty frustules on the northern transect suggested a generally declining population. However, Phaeocystis sp. was more abundant on the northern transect.     

Diatoms of the microphytobenthic community in a tropical intertidal sand flat influenced by monsoons: spatial and temporal variations

Seasonal variations in the microphytobenthic diatom community were investigated in an intertidal sand flat of a tropical marine environment influenced by monsoons. Cores of sediments were collected along the beach gradient: low tide, mid tide and high tide zone up to a depth of 15 cm.. Diatom abundance was lowest during the monsoons and highest during the post-monsoons and the early pre-monsoon season throughout the intertidal transect. Diatom diversity was highest at the mid tide, followed by the high and low tide zones. Diatoms were viable up to a depth of 15 cm throughout the intertidal transect. The diatom community included the pennates, the permanent residents of this area, centric genera, which lead an attached mode of life and also some planktonic genera, brought in from ambient waters. Among the pennates, Navicula and Amphora were the dominant genera whereas in the case of centrics, Thalassiosira dominated the community throughout the intertidal transect down to a 15 cm depth. . Grain size fractions, which served as predictors of some diatom genera changed with tidal zones. The effect of winds on the resuspension of the pennate diatoms was evident only at the low tide zone down to a depth of 5 cm . Chlorophyll a concentration proved to be a good predictor of both pennate and centric diatom abundance at the low tide zone down to a depth of 10 cm and at the mid tide zone down to a depth of 5 cm.. However, even though chlorophyll a concentrations failed to reveal any positive correlation with the diatom abundance at both the deeper sediment layers and the high tide zone, the fact that viable cells were present at these areas reveal that the diatoms adopt survival strategies, contributing significantly to the carbon budgets of such unstable habitats.     

Saturated uptake kinetics: transient response of the marine diatom Thalassiosira pseudonana to ammonium,nitrate, silicate or phosphate starvation

Changes in the saturated uptake kinetics of the limiting nutrient were followed as Thalassiosira pseudonana (Clone 3 H) batch cultures entered ammonium, nitrate, silicate and phosphate starvation. Cultures starved of ammonium or phosphate developed very high specific uptake capacities over a 24 to 48 h starvation period, due to both decreases in cell quota and increases in uptake rates per cell. In particular, the cell phosphorus quota decreased ca. 8-fold during phosphate starvation and specific uptake rates exceeded 100 d^-1. In contrast, cultures entering nitrate or silicate starvation underwent little or no further cell division, and the uptake capacity declined during starvation. After 24 to 48 h starvation, an induction requirement for uptake of nitrate or silicate was apparent. These responses are consistent with adaptation to the pattern of supply of these nutrients in the field.     

Spring bloom sedimentation in a subarctic ecosystem

Results from a 5-yr study (1985 to 1989) in Auke Bay, Alaska show that termination of the spring bloom consistently occurred at limiting nitrate concentrations. Following nutrient exhaustion, phytoplankton sinking rates increased and displayed greater temporal variability. Threshold nitrate concentrations, approximating K_s values of the species present, were found to signal initiation of increased sedimentation. For Thalassiosira aestivalis, the threshold was ～2 μmol l^-1, while for Skeletonema costatum the threshold was ～1 μmol l^-1, suggesting genus-specific differences in sinking-rate sensitivity to nitrate exhaustion. Overall, sinking rates of the three principal genera ranked (high to low) Thalassiosira spp.> S. costatum>Chaetoceros spp., while the nitrate sensitivities of the sinking rates of the genera ranked (high to low) Thalassiosira spp.> Chaetoceros spp.> S. costatum. Thalassiosira spp. showed the most consistent sinking rate increases following nutrient impoverishment. During a bloom dominated by T. aestivalis, a decrease of cell sinking rate with depth coincided with a decrease in short-term nutrient stress as measured by intracellular nitrate pools. In addition, no correlation was found between chain length or aggregate formation and sinking rate for this species. Though we measured only small-scale cell-cell adhesion, not larger-scale marine snow formation, this supports the notion that the sinking rates of Thalassiosira spp. were controlled primarily by cell physiology. For S. costatum, however, shorter chains sank faster. The sinking behavior of the species studied here figures prominently in their pelagic ecology and in the carbon flux of coastal ecosystems, both of which are driven by short-term variability.     

A diatom assemblage living below the surface of intertidal sand flats

The assemblages of species living on and in intertidal sand flats at Barnstable Harbor, Massachusetts (USA) are briefly described. An endopsammic assemblage of diatoms, mainly Amphora species, living below the surface is described for the first time in such a marine situation. Some aspects of its ecology and relationship to the other diatom assemblages living on the sediments are discussed.     

Potential for increased nutrient uptake by flocculating diatoms

Blooms of chain-forming diatoms commonly flocculate into centimeter-sized aggregates of living, vegetative cells following nutrient depletion in surface waters off southern California. We examined the hypothesis that diatom cells within aggregates experience increased nutrient uptake relative to unattached cells. We measured in situ settling velocities of 49 to 190 m d^–1 and calculated porosities of 0.99931 to 0.99984 (±>0.03%) for 12, newly-formed diatom flocs ranging from 0.19 to 4.2 cm³ in volume and 7 to 22 mm in equivalent spherical diameter. Using permeability-porosity relationships, we calculated intra-aggregate flow velocities of 20 to 160 m s^–1. Although subject to considerable uncertainty, a Relative Uptake Factor analysis based on mass transfer equqtions suggests that diatoms fixed within aggregates undergoing gravitational settling can take up nutrients up to 2.1±0.4 times faster than unattached diatoms experiencing laminar shear. Increased nutrient uptake by aggregated diatoms may be importan in understanding the reasons for diatom floc formation.     

A new source of dimethylsulfide (DMS) for the arctic atmosphere: ice diatoms

We report the first evidence that pennate diatoms growing within the bottom layer of first-year ice in the Arctic produce significant amounts of particulate dimethylsulfoniopropionate (DMSP_p) and dissolved DMSP+DMS. In 1992 in Resolute Passage, a tributary of Barrow Strait, DMSP_p concentrations within the bottom layer of ice reached 1055 mg S m^-3 at the end of the vernal bloom, a value one order of magnitude higher than the maximum value reported in antarctic ice. Bottom-ice concentrations in DMSP_p and DMSP_d+DMS were significantly correlated with the abundance of the dominant pennate diatom Nitzschia frigida. Intracellular concentration in DMSP of ice algae was very low (0.001 pg cell^-1) at the end of April when algae were light-limited and reached 1.17 pg cell^-1 in mid-May following an increase in light and algal growth. We calculate that the rapid release of the dissolved DMSP+DMS from the ice into surface waters following the ice break-up will generate a sea-to-air DMS flux of 0.7 mg S m^-2 d^-1, a pulse ten times higher than the mean arctic summer flux. We estimate that this 1-d pulse represents up to 5% of the annual DMS emission in the Arctic.     

Effects of silicate depletion on photosynthesis by diatoms in the plume of the Hudson River

Silicate depletion was observed during a bloom of netplankton diatoms. Netplankton chlorophyll a increased over the same salinity range and at the same rate that silicate decreased. Silicate depletion coincided with a decrease in the apparent Si:N uptake ratio as well as a change in the magnitude and diurnal phasing of light saturated photosynthesis (P _m ^B ) by netplankton diatoms. Nanoplankton P _m ^B was unaffected by silicate depletion and increased with temperature. Consequently, nanoplankton P _m ^B eventually exceeded netplankton P _m ^B while netplankton biomass was still increasing relative to nanoplankton biomass.     

Nitrate- and silicate-competition among antarctic phytoplankton

Natural phytoplankton from antarctic waters in the Drake Passage were used for competition experiments in semicontinuous cultures. The outcome of interspecific competition for silicate and nitrate was studied at a range of Si:N ratios (from 2.6:1 to 425:1) and at three different dilution rates. For five species Monod kinetics of silicate-and nitrate-limited growth has been established. Comparison between theoretical predictions derived from Monod kinetics and the outcome of competition experiments showed only minor deviations. Contrary to literature data, considerable depletion of nitrate was found in antarctic seawater. Both the concentrations of soluble silicate and of nitrate were too low to support maximum growth rates of some of the diatom species under investigation.     

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.     

Hydrodynamic and chemical conditions during onset of a red-tide assemblage in an estuarine upwelling ecosystem

The hydrodynamics and nitrogen/silicon biogeochemistry accompanying the development of a red-tide assemblage were examined in the Ría de Vigo (northwest Spain), a coastal embayment affected by upwelling, during an in situ diel experiment in September 1991. Despite a low N:Si molar ratio (0.5) of nutrients entering the surface layer, which was favourable for diatom growth, the diatom population began to decline. Limited N-nutrient input, arising from moderate coastal upwelling in a stratified water column, restricted net community production (NCP = 630 mg C m⁻² d⁻¹). In addition, light-limitation of gross primary production (GPP = 1525 mg C m⁻² d⁻¹) was observed. The relatively high f-ratio (= NCP:GPP) recorded (0.41, characteristic of intense upwelling conditions) would have been as low as 0.15 had not GPP been limited by light intensity. Temporal separation of carbohydrate synthesis during the photoperiod from protein synthesis in the dark could be inferred from the time-course of the C:N ratio of particulate organic matter. Severe light-limitation would lead to diatom collapse were the diatoms not able to meet all their energy requirements during the hours of darkness. Under the hydrodynamic, nutrient and light conditions of the experiment, an assemblage of red-tide-forming species began to develop, aided by their ability to migrate vertically and to synthesize carbohydrates during the light in surface waters and protein during the dark at the 4 m-deep pycnocline. Thermal stratification, reduced turbulence, intense nutrient mineralization, and the limited nitrogen input through moderate upwelling were all favourable to the onset of a red-tide assemblage. Received: 15 February 1997 / Accepted: 26 September 1997     

Feeding habits of marine cladocerans in the Inland Sea of Japan

Natural food items of five species of marine cladocerans, Evadne nordmanni, E. tergestina, Penilia avirostris, Podon leuckarti and P. polyphemoides, were investigated in the Inland Sea of Japan between April 1986 and May 1987. Gut content examination with SEM (scanning electron microscopy) revealed that feeding was largely limited to centric diatoms and a few exceptions of pennate diatoms and dinoflagellates. No animal remains were detected, and some unidentified materials were also found. Phytoplankton smaller than 35 m in size (cell diameter in centric diatoms and longest dimension in others) was found most frequently in the gut of cladocerans. The role of grazing of marine cladocerans in trophodynamic pathways of the pelagic realm is discussed.     

Simultaneous nitrogen and silicate deficiency of a phytoplankton community in a coastal jet-front

The nutritional status of a phytoplankton community was investigated in a coastal jet-front located in the Gulf of St. Lawrence, Canada, in 1987. During the sampling period, the frontal community was mainly composed of the diatomsChaetoceros debilis, Skeletonema costatum, Thalassiosira gravida andC. pelagicus. As previously reported for the St. Lawrence, some frontal stations were depleted both in nitrate and silicate. At stations impoverished in nitrate, internal nitrate pool concentrations were low or undetectable, suggesting that cells had not, recently, been exposed to a nitrate flux which exceeded the nitrate assimilation rate. At these impoverished stations, however, ambient and intracellular concentrations of ammonium and urea were high, suggesting that the community was not nitrogen-deficient. The comparison between the ambient silicate concentrations and the silicate requirement (K _s ) of the dominant diatoms suggests thatC. debilis andS. costatum were Si-deficient. This is further supported by the low silicate uptake rates and intracellular concentrations measured at the silicate impoverished stations. The silicate deficiency also resulted in a decrease in the seston and phytoplankton N:C ratios.Please address all correspondence and requests for reprints to Dr Levasseur at his new address: Maurice Lamontagne Institute, 850 Route de la Mer, Mont-Joli, Québec G5H 3Z4, Canada     

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.     

Phytoplankton sinking rates in oligotrophic waters off Hawaii,USA

The sinking rates in two size fractions of natural phytoplankton were measured over much of the photic zone in a subtropical environment. At 24, 40 and 71 m, sinking rates (± SD) of the 3 to 20 m fractions were 0.72±0.05, 0.83±0.05, and 0.34±0.04 m · d^-1, respectively, and rates for the 20 to 102 m fraction were 1.50±0.21, 1.65±0.14, and 0.95±0.22 m · d^-1, respectively. At all depths sampled, the 20 to 102 m size fraction was observed to sink significantly (P<0.01) faster than the 3 to 20 m fraction. considering vertical variability, both size fractions were observed to have significantly (P<0.01) lower sinking rates at 71 m than at more shallow depths. The finding of lower sinking rates in the region which lies just above the subsurface chlorophyll maximum provides empirical support for the hypothesis that variations in phytoplankton buoyancy may be related to the maintenance of this feature which is typical in oceanic environments. Analysis of sinking rate traces describes the distributions of specific sinking velocities for each size fraction and their variation.Oceanic Institute Contribution No. 162