Dynamique du phytoplancton et matière organique particulaire dans la zone euphotique de l'Atlantique Equatorial

At two fixed stations in the Equatorial Atlantic Ocean (0°–4° W), the physical, chemical and biological properties of the euphotic layer were determined for 14 d (Station A: 5–18 February, 1979) and 13 d (Station B: 20 October–7 November, 1979), respectively. The stability of the water column allowed comparison of 3 different “systems”: (i) a well-illuminated and nitrate-depleted mixed layer; (ii) a chlorophyll maximum layer (chl a _max) in the thermocline which is poorly illuminated (6.3% of surface irradiance); (iii) a well-illuminated but nitrate-rich (>0.9 μg-at l^-1) mixed layer. In each layer the particulate organic carbon (COP), nitrogen (NOP) and phosphorus (POP) contents were measured and compared with the phytoplankton biomass. In the chlorophyll maximum layer, the phytoplankton biomass contributed significantly to the total particulate organic matter (between 55 and 75%). In the nitrate-depleted mixed layer, the results varied according to whether the regression technique [COP=f(chl a)] was used, or the chl a synthesis during the incubation of the samples. With the former technique, the phytoplankton carbon (C _p) content appeared minimal, because the y intercept, computed using all the data of the water column, was probably overestimated for this layer. POP would be more associated with living protoplasm than with carbon and nitrogen in the three layers. In the chlorophyll a maximum layer it constitutes a valuable detritus-free biomass measurement, since 80% of the POP consist of phytoplankton phosphorus. The assimilation numbers (NA=μg C μg chl a ^-1 h^-1) were high in all three layers, but the highest values were recorded in the nitrate-depleted mixed layer (NA=15 μg C μg chl a ^-1 h^-1). In the chlorophyll maximum layer, light would be a limiting factor during incubation: between 10²⁵ and 8.10²⁴ quanta m^-2 d^-1 NA and light are positively correlated independant of nitrate concentration. The growth rates of phytoplankton (μ) were estimated and compared to the maximum expected growth rate. Our main conclusion was that despite very low biomass and nutrient content, the mixed layer was in a highly dynamic state, as evidenced by high rates of phytoplankton growth and short nutrient turnover times (1 d or less for PO-P₄ in the mixed layer versus 3 d in the thermocline). The presence of nitrate in the water column allows the development of a higher phytoplankton biomass but does not increase growth rate.     

In-situ measurements of nitrogenous nutrient uptake by kelp (Ecklonia maxima) and phytoplankton in a nitrate-rich upwelling environment

Nitrogen uptake by the kelp Ecklonia maxima Osbeck and phytoplankton was examined under different conditions of nutrient availability in a kelp bed off the Cape of Good Hope by measuring nutrient depletion in large plastic bags by the kelp and ¹⁵N uptake by phytoplankton. E. maxima took up nitrate and ammonia, but not urea, and showed only a weak preference for reduced nitrogen. Phytoplankton absorbed all three forms of nitrogen available, with a preference for ammonia and urea. Ambient nitrate concentration exhibited a marked and rapid decrease with northerly winds and an increase in response to offshore southerly winds. Nitrogen uptake by E. maxima was linearly related to ambient concentration and did not saturate even at nitrate concentrations >20g-at N l^-1, resulting in a significantly higher tissue nitrogen content under upwelling conditions. Nitrate imported by upwelling was the chief source of nitrogen utilised within the kelp bed. Locally regenerated nitrogen (ammonia and urea) was calculated to contribute only ca 4% of total nitrogen uptake during upwelling and 30% during the relaxation or downwelling phase.     

Nitrate storage by phytoplankton in a coastal upwelling environment

The storage of nitrate by phytoplankton cells during the early phases of upwelling was studied in coastal stations off northern Spain (southern Bay of Biscay) between 1990 and 1994. In this region, a persistent upwelling during summer is characterised by intermittent pulses of variable intensity, and increased nutrient concentrations in the surface layer. The main effect of an upwelling pulse on phytoplankton distribution is the shifting of the chlorophyll a and primary production maxima to near the surface. When the upwelling relaxes, thermal stratification of the water column occurs, and a distinct subsurface chlorophyll maximum develops below the production maximum. An accumulation of intracellular nitrate characterized the early phases of upwelling (mean = 2.73 μmol N m⁻³), maximum concentrations being attained at depths where biomass and production values were moderate. In contrast, phytoplankton cells from non-upwelling situations contained significantly lower concentrations of intracellular nitrate (mean = 0.17 μmol N m⁻³). The variations in the intracellular pool of nitrate may result from the differential allocation of resources within the cell as a result of variations in the energy available, since the uptake and assimilation of nitrate is a relatively expensive process involving several enzymatic systems. We hypothesize that nitrate storage by phytoplankton cells is characteristic of early phases of upwelling and is linked to patterns of carbon fixation. Average nitrogen budgets for upwelling and non-upwelling situations indicate that intracellular nitrate reserves are not responsible for maintaining high phytoplankton growth rates, since they only account for <2% of daily primary production during upwelling events. Received: 28 August 1996 / Accepted 3 December 1996     

Nitrate metabolism in abyssal waters

Activities of nitrate assimilation and nitrate reduction were measured 50 cm above the ocean floor (5,845 m and 5,207 m) by an in situ ¹⁵N tracer technique at stations in the subtropical (28°29.8′N; 144°58′E) and subarctic (44°10.2′N; 154°03′E) western North Pacific Ocean. Nitrate assimilation ranged from 0.009 to 0.11 μg-at N/1/day, and nitrate reduction from 0 to 0.42 μg-at N/1/day in the presence of added peptone and yeast extract. Nitrate assimilation was higher than nitrite formation at the southern station, but the reverse was the case at the subarctic station. No correlation was observed between bacterial growth and nitrate metabolizing activities. Data are also presented on the effect of hydrostatic pressures upon nitrate metabolism by microbial populations in the surface waters.     

Coccolithophores: a major phytoplankton component in mature upwelled waters off the Cape Peninsula,South Africa in March, 1983

Coccolithophores, together with other phytoplankton, were sampled at ten stations along a transect following the general orientation of the Cape Peninsula upwelling plume in March, 1983. Thirteen coccolithophore species were identified with Emiliania huxleyi dominating the assemblages. Coccolithophores and other microflagellates were more abundant than diatoms along the transect except at one station in mature upwelled water. Very high populations (2 340×10³ cells l^-1) of coccolithophores were found in mature upwelled water adjacent to the oceanic front. This count is one of the highest counts ever recorded in an upwelling area. The anomalous warm event prior to sampling may have had a residual effect on seeding, contributing towards the predominance of small-celled phytoplankton over chain-forming diatoms.     

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.     

Distribution of chlorophyll,carotenoids and phytoplankton in relation to certain environmental factors along the central west coast of India

Distribution of chlorophyll pigments, carotenoids and abundance of phytoplankton in relation to certain environmental factors of the nearshore waters off the central west coast of India (latitudes 15°30 to 18°30N) were studied monthly at 7 stations during 1970/1971. Changes in the hydrographical factors and the biological processes occurring in the region during different months appear to be influenced by the pattern of upwelling along the northern and southern parts of the west coast of India. The pigment concentration shows a marked decrease in October, but is followed by a slow but steady rise, which reaches its maximum in April/May. A slightly smaller maximum is noticed in December/January. The composition of various chlorophyll pigments and carotenoids indicated the physiological state of phytoplankton populations during different months in the region investigated. Abundance of specific phytoplanktonic elements, consisting mainly of diatoms, in space and time, characterises the waters of the central west coast of India, indicating a clear succession of species.     

Variability of biochemical composition and size distributions of seston in the euphotic zone of the Bay of Biscay: implications for microplankton trophic structure

Particle-size distributions and several biochemical components of seston were studied from October 1986 to December 1987 in surface waters of the Bay of Biscay. Variance partitioning of hydrographic and seston parameters indicates that, although seasonal variability related to the thermal cycle was important, most of the differences in seston concentration were caused by spatial components of variance. Changes in the vertical structure of the water column appeared to be the principal source of variability. Phytoplankton blooms in spring and fall and the effects of the coastal upwelling and the thermocline during summer were traced using seston concentration and biochemical composition. The different seston measurements were scaled according to their correlations with total concentration and particle size. Photosynthetic pigment biomass was related to both large and small partieles. However, concentrations of particulate proteins, lipids and carbohydrates were more related to small particles. Although not specifically analysed, low concentrations of inorganic particles and detritus can be expected in the study area, thus most of the particulate organic matter was associated with small, non-pigmented organisms, particularly in surface waters during the period of thermal stratification. The variability in concentrations of total seston and particulate organic matter was mainly due to variations in particulate proteins, lipids and carbohydrates; pigment concentrations were of secondary importance. Expressed as the ratio chlorophyll a: particulate protein-nitrogen, phytoplankton constituted an important fraction of the microplankton biomass only during spring blooms, when it averaged 75% of the particulate protein-nitrogen. In constrast, <30% of protein-nitrogen was related to chlorophyll a during the summer. These results suggest that an oligotrophic structure based on the microbial-loop prevails in microplankton assemblages of surface waters for most of the year.     

Significance of cellular nitrate content in natural populations of marine phytoplankton growing in shipboard cultures

Phytoplankton intracellular nitrate concentrations have been monitored in a 56-h experiment on a shipboard culture of surface sea water from an upwelling region. These measurements were related to parameters of biomass (particulate nitrogen) and nitrate assimilation using the ¹⁵N isotope technique and the nitrate reducase (NR) assay. The procedure for measuring cellular nitrate concentrations is described. This parameter exhibited diurnal variations, ranging from 3.1 to 20.6 ng-at nitrate per g-at particulate nitrogen, and could be correlated positively with NR activity. Nitrogen budgets show that NR activity represents only 12% of nitrate incorporation in organic phytoplankton material when nitrate is available in the sea water. However, upon depletion of the environmental nitrate (zero uptake), NR activity can fully account for the decrease of internal nitrate. From the results, it seems that internal nitrate content is a better index of nitrate consumption by marine phytoplankton than the external concentration of nitrate-nitrogen.     

Variability of particulate organic carbon and nitrogen in the Namibian upwelling system

Particulate organic carbon (POC) and nitrogen (PON) were analyzed during cruises undertaken in September 1985 and April 1986 along the Namibian upwelling system. The main objectives were to provide estimates of standing stocks of particulate organic matter (POM) and analyze its temporal and spatial variability. Average estimates of total carbon standing stock (0 to 100 m depth) accounted for 1.2 g-at C m^–2 during active and 1.32 g-at C m^–2 during abated upwelling. Zooplankton carbon was estimated as 0.22 and 0.27 g-at C m^–2, respectively, for both periods. POM was generally concentrated near the surface, especially during abated upwelling. During abated upwelling, POM was not only abundant inshore but also over the shelf, as a response of a more diffuse frontal system and a more strongly stratified water column. Cross-shelf gradients were more significant during active upwelling, while alongshore gradients accounted for most of the variance of particulate organic matter during abated periods. This result was a consequence of the seasonal intrusion of warm, Angolan water from the north during the period of minimum upwelling, and resulted in poorer POM concentrations and higher consumer: producer ratios (24.4%). Nevertheless, this last conclusion should be regarded with caution due to the lack of comparative interannual variability. A 48 h study at a fixed station permitted analysis of the daily variability in POM during the intrusion process. Changes in the thickness of the surface mixed layer due to irregular time-spaced pulses of non-homogeneous water masses resulted in sudden enrichments and renewals of phytoplankton and zooplankton populations in a matter of hours.     

Primary production and environmental factors in an oligotrophic biome in the Aegean Sea

The rate of the primary production of the phytoplankton community in the Petalion Gulf, Aegean Sea, was studied from January 1970 to May 1971, at a station situated at approximately Latitude 37°54N; Longitude 24°11E. A variety of physical and chemical parameters such as chlorophyll, primary nutrients (N,P,Si), temperature, salinity, oxygen and light penetration were also studied simultaneously. The rate of the gross primary production varied from 40 to 200 mg C m^-2 day^-1, with a mean value of 90 mg C m^-2 day^-1. The annual gross primary production was calculated to be 33 g C m^-2, which is the minimum known value in the Aegean and Mediterranean Seas. Maximum production was found at the depth of 20 m on the average, mainly due to high light intensities. Petalion Gulf supports a small photosynthetic biomass, as indicated by the low seasonal values of chlorophyll a (0.01 to 0.18 mg m^-2), the highest values being found in the summer. The low production rate noted may have been due to the low nutrient concentrations found: N, 0.04 to 0.32 g-at/1; P, 0.00 to 0.15 g-at/1; Si, 0.45 to 2.25 g-at/1. It is suggested that inorganic phosphorus and nitrogen may alternate in limiting primary production rates in these oligotrophic waters. The temperate waters of the Petalion Gulf are stratified in summer (15.5° to 24.7°C) and well-mixed in winter (12.9° to 15.0°C); they are oxygen-saturated throughout the year, and of high transparency, with 86 m depth for the euphotic zone on the average yearly. The Petalion Gulf is therefore characterized as a typical oligotrophic biome in the Aegean and Eastern Mediterranean Seas.     

Nitrogen assimilation by phytoplankton and other microorganisms in the surface waters of the central North Pacific Ocean

Assimilation rates of ¹⁵N-labelled ammonium, urea, and nitrate by plankton in the upper euphotic zone were measured in 2 summer, 2 winter, and 1 spring cruise in the central North Pacific Ocean. Average rates of ammonium plus urea assimilation could not be determined precisely, but were estimated to be 7 to 25 g-at. N m^-3 day^-1. Indirect evidence suggested that non-photosynthetic microorganisms contributed to these rates. Nitrate assimilation was negligible in the upper waters considered in this report (above the chlorophyll maximum and the nutricline). Potential, nitrate-saturated rates were in the range 1 to 8 g-at. N m^-3 day^-1. Seasonal comparison showed lowest rates of both carbon and nitrogen assimilation rates per chlorophyll a in winter.     

黄海春季表层叶绿素和初级生产力及其粒径结构研究

根据2006年4月对黄海浮游植物分级叶绿素及初级生产力的调查,研究了黄海叶绿素及初级生产力的水平分布及粒级结构特征,并分析了其主要影响因素。黄海海域调查站位表层叶绿素a质量浓度变化范围为0.20～4.94μg·L-1,平均值为0.96μg·L-1。叶绿素最大值出现在临近长江口的站位。叶绿素分级结果表明黄海春季以粒径〉5μm的浮游植物占优势。黄海表层初级生产力的变化范围为2.03～15.64mg·m-3·h-1,平均值为6.08mg·m-3·h-1。其中南黄海海域初级生产力平均为6.58mg·m-3·h-1,北黄海海域初级生产力平均为4.92mg·m-3·h-1。高值区分布在南黄海中部。受水体透明度的影响,低值区出现在临近长江口的站位。断面站位分析表明浮游植物初级生产力由北向南逐步升高,温度随纬度的变化是南北海域初级生产力水平差异的主要原因。由于粒径较小（〈5μm）的浮游植物单位叶绿素具有较高的碳固定能力,调查期间整个海区初级生产力以粒径〈5μm的浮游植物贡献为主。     

Phytoplankton biomass and production in the upwelling region of NW Africa. Relationships with hydrographic parameters

Phytoplankton biomass and production in the upwelling region of NW Africa and relationships with hydrographic parameters were studied. During the cruise of Atlor VII carried out in November 1975 in the upwelling region of NW Africa, measurements of chlorophyll a and primary production as derived from ¹⁴C uptake experiments were made at a total of 40 stations. Biomass and production showed the higher values on the shelf in the area of Banc d'Arguin and north of Cap Blanc. Production estimates in this area ranged between 1.4 and 3.2 g C m^-2 d^-1. There was a marked minimum in biomass and production at Lat. 21 ° N, in the zone of maximum upwelling intensity. With the exception of this minimum, the productive area coincided with the zone where surface temperature was lower than 18 ° C (indicating dominance of upwelled Central Waters) and nutrients were detectable in the upper layers. In the poorer offshore area there was a distinct subsurface chlorophyll maximum. The results are compared with those of previous cruises and some features of the seasonal cycle in the studied area are discussed.     

Eddy-induced oscillations of the pycnocline affect the floristic composition and depth distribution of phytoplankton in the subtropical Pacific

Changes in the phytoplankton population caused by a wind-induced mesoscale eddy, first located off the Hawaiian chain on 29 August 1989, were examined from 6 to 12 September 1989. These cyclonic eddies produce upwelling of nutrient-rich water into the photic zone which may induce changes in the distribution of phytoplankton. The eddy affected the depth, magnitude and composition of the chlorophyll maximum. The eddy produced an upward shift in the distribution of phytoplankton, shoaling the chlorophyll maximum to 30 m. Pigment concentrations at the chlorophyll maximum were enriched up to 2.2 times inside the eddy. Outside the eddy, the upper mixed layer (UML, 50 m) was dominated by cyanobacteria whereas the deep chlorophyll maximum (DCM) was dominated by prochlorophytes, chlorophytes and chrysophytes. Inside the eddy, the UML showed an increase in deep water taxa but little change in cyanobacteria. The observed population changes probably resulted from increased growth rates and vertical transport of cells. Based on projections of the track of the eddy, we inferred that its trail created an area of enhanced chlorophyll that was not quickly consumed, and concluded that coupling between phytoplankton and grazers was weak. The presence of such cyclonic eddies may explain some of the variability in chlorophyll concentration found in the oligotrophic subtropical Pacific.     

Interactions of inorganic nitrogen in the uptake and assimilation by marine phytoplankton

The effect of ambient ammonium concentration on the nitrate uptake rate of marine phytoplankton was investigated. These studies consisted of laboratory experiments using unialgal species and field experiments using natural phytoplankton communities. In laboratory experiments, ammonium suppressed the uptake rates of nitrate and nitrite. Approximately 30 min were required for ammonium to exhibit its fully inhibitory effect on nitrate uptake. At high ammonium concentration (>3 g-at/l), a residual nitrate uptake rate of approximately 0.006 h^-1 was observed. When the ambient ammonium concentration was reduced to a value less than 1 g-at/l, the suppressed nitrate uptake rate subsequently attained a value comparable to that observed before the addition of ammonium. A range of 25 to 60% reduction in the nitrate uptake rate of natural phytoplankton communities was observed at ambient ammonium concentrations of 1.0 g-at/l. A mechanism is proposed for the suppression of nitrate uptake rate by ammonium through feedback control of the nitrate permease system and/or the nitrate reductase enzyme system. The feedback control is postulated to be regulated by the level of total amino acids in the cell.Contribution No. 936 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA. This paper represents a portion of a dissertation submitted to the Department of Oceanography, University of Washington, Seattle, in partial fulfillment of the requirements for the Ph.D. degree.     

Diel oscillations of the photosynthesis-irradiance (P-I) relationship in natural assemblages of phytoplankton

Diel oscillations in the photosynthesis-irradiance (P-I) relationship are described for marine phytoplankton assemblages at 6 stations in an upwelling area off the southern California coast (USA) between May and August 1980. The initial slope () and asymptote (P _max) of P-I curves changed significantly over the day; both parameters were in phase and had similar changes in amplitude. The diel oscillations in photosynthesis appeared unrelated to changes in chlorophyll a concentrations. Amplitudes of daily variations in photosynthesis ranged from approximately 3 to 9, as measured by the maximum to minimum ratio for photosynthetic capacity (P _max). Diatom-rich samples collected during an upwelling event and those dominated by dinoflagellates both had midday to early afternoon maxima in and P _max. Samples from other locations had peak photosynthetic activity later in the afternoon. The relationship between and P _max was consistent in all phytoplankton samples analyzed, with a surprisingly high correlation considering the spatial and temporal scales encompassed in this study. These results indicate that the photosynthesis-irradiance (P-I) relationship is time-dependent and, moreover, that changes in and P _max are closely coupled for a variety of natural phytoplankton assemblages.     

Ecotypic differentiation of Laminaria longicruris in relation to seawater nitrate concentration

The ultraplankton (cell diameters >3 μm), which compromises about 70% of the biomass of phytoplankton in subtropical surface waters near Oahu, Hawaii, was isolated for growth rate studies. The specific growth rate (μ) was estimated from the rate of increase of the chlorophyll biomass during incubations in the absence of grazers. This growth rate of the ultraplankton ranged from 0.037 to 0.071 h^?1 (=1.3 to 2.5 doublings d^?1) during a period when P:B ratios of 5 to 14.5 μg C μg^?1 chl a h^?1 prevailed. The co-occurrence of atypically high P:B ratios and nonlimiting ambient nutrient concentrations suggests that the calculated values are higher than those characteristic of such subtropical ecosystems in general. Rates of ammonium uptake and photosynthesis by the >3 μm fraction were also compared to those of larger fractions. Organisms in the >3 μm fraction assimilated NH ₄ ⁺ at a rate which was about 75% greater than that of the 3 to 20 μm size fraction. Comparison of μ and P:B data collected over a 2 mo period (November–December, 1980) shows that the correlation between these two rate indices is nonlinear. The predominance of small-celled phytoplankton in oligotrophic waters is explained, in part, by its higher μ, its higher nutrient assimilation rates, and the absence of its loss through sedimentation.     

Light acclimation states of phytoplankton in the Southern Ocean, determined using photosynthetic pigment distribution

In high-latitude waters such as the Southern Ocean, the primary production of phytoplankton supports the ecosystem. To understand the photo-acclimation strategy of such phytoplankton within cold environments, the vertical distribution profile of photosynthetic pigments was analyzed in the Southern Ocean. Samples were taken along 110°E during the austral summer, and along 150°E and around the edge of the seasonal sea ice of the Antarctic Continent during the austral autumn. Pigment extraction methods were optimized for these samples. The standing crop of chlorophyll a was larger in the region along the edge of the seasonal sea ice than at sampling stations in open ocean areas. Chlorophyll concentration seemed to be dependent on the formation of thermo- and haloclines along the edge of the seasonal sea ice, but not in the open ocean where such clines are less pronounced. The marker pigments fucoxanthin and/or 19′-hexanoyloxyfucoxanthin were dominant at most sampling stations throughout the water column, while other marker pigments such as alloxanthin were quite low. This indicated that diatoms and/or haptophytes were the major phytoplankton in this area. Comparison of the relative ratio of fucoxanthin with that of 19′-hexanoyloxyfucoxanthin allowed some stations to be characterized as either diatom-dominant or haptophyte-dominant. The relative ratio of xanthophyll-cycle pigments (diadinoxanthin plus diatoxanthin) to chlorophyll a was high in surface waters and decreased gradually with depth. This suggests that near the ice edge during summer in the Southern Ocean, both diatoms and haptophytes acclimate to their light environments to protect their photosystems under high-light conditions.     

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

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