Comparison of bacterial and algal utilization of orthophosphate in an estuarine environment

Bacterial utilization of orthophosphate in an estuarine environment has been differentiated from algal utilization by using flow-filters of 5.0, 1.2 and 0.45 m poresize. Examination by light microscopy showed that most of the bacterial population passed through a 5.0-m filter, whereas most algae were retained. In all experiments, bacterial and algal cell numbers and biomass were estimated. P-uptake by algae and bacteria was closely correlated with cell biomass. P-uptake by algae was high only in the summer months, whereas P-uptake by bacteria was high throughout the year. Neither algal nor bacterial P-uptake, however, was correlated with temperature or dissolved orthophosphate, total organic phosphate or total phosphate concentrations. Cell biomass of algae at a given time had a high correlation with dissolved organic phosphate levels in 2 weeks prior to sampling (r=0.830) and a low correlation in the 2 weeks following sampling (r=0.0005). Algal cell numbers had a high correlation with bacterial cell numbers (r=0.950). The biomass of algae and bacteria also had a high correlation (r=0.902). The rate of P-uptake from the water by algae and bacteria varied with season and with the species composition of the natural population.     

Diet of Neotropic cormorant (Phalacrocorax brasilianus) in an estuarine environment

The diet of the Neotropic cormorant (Phalacrocorax brasilianus) was studied by analysing 289 regurgitated pellets collected from a roosting site at Lagoa dos Patos estuary, southern Brazil, between November 2001 and October 2002 (except April to June). In total, 5,584 remains of prey items from 20 food types were found. Fish composed the bulk of the diet representing 99.9% by mass and 99.7% by number. The main food items were White croaker (Micropogonias furnieri) (73.7% by frequency of occurrence, 48.9% by mass and 41.2% by number), followed by Catfish (Ariidae) and anchovies (Engraulididae). In Lagoa dos Patos estuary the generalist Neotropic cormorant fed mainly on the two most abundant demersal fishes (White croaker and Catfish), which accounted for the low niche breadth calculated. The total length of all fish preyed varied from 27.2 to 318.3 mm (113.5 ± 48.0 mm), and preyed White croakers’ size differed between months. Neotropic cormorants seem to prey on most abundant class sizes of White croaker instead of selecting similar prey size throughout the time. However, temporary changes in diet in terms of food items, abundance and prey size were detected, revealing a high ecological plasticity of the species. Individual daily food intake of Neotropic cormorants estimated by pellets and metabolic equations corresponded to 23.7 and 27.1% of their body mass, falling in the range of other cormorant species. Annual food consumption of the population estimated by both methods was 73.4 and 81.9 tonnes, comprising mainly immature and subadult White croaker and Catfish which are commercially important. Temporal variations in diet composition and fish size preyed by Neotropics cormorants, a widespread and generalist species, suggest shifts according to fluctuations in the abundance of prey. The plasticity of this cormorant is also revealed by their ability to adjust feeding behaviour in response to temporal or local changes in the environment, from a generalist at the species level to a specialist at the individual or local population level.     

Seasonal,geographical and size-induced variability in mercury content of Mytilus edulis in an estuarine environment: a re-assessment of mercury pollution level in the Estuary and Gulf of St. Lawrence

Seasonal variability of mercury content of Mytilus edulis (L.) was studied at monthly intervals for 18 mo (June 1978–December 1979) in a natural population from a typical estuarine environment (St. Lawrence Estuary). Body-size-induced variability was observed in the same population in spring 1980. The regression coefficients of log-log relationships between mercury content and body size of the mussels varied with time. However, the regression coefficients were never higher than 1; this indicates, therefore, no net accumulation of mercury with age of the bivalve. A normalization method, taking into account shell length and soft-tissue dry weight, is proposed to minimize the metal variations due to size differences of the mussels. The normalized mercury contents vary seasonally by a factor of two. Minimum values were observed during summer. This factor (2) is taken as the minimal difference which can be accepted as indicating a significant geographical trend in a mussel watch-type study in estuarine environments. In the St. Lawrence Estuary and Gulf, size-normalized concentrations range from 69 to 378 ng Hg g^-1 (dry weight); highest levels were observed in areas where freshwater influence is maximum, while lower contents were encountered in regions of highest Atlantic influence. In the Gulf, mercury in mussel soft tissue is 3x10⁴ times that of the seawater. There is some indication of a decrease in the previous measured levels near the mouth of the Saguenay fjord in the Lower St. Lawrence estuary.     

Respiratory responses of the estuarine mysid Neomysis integer (Peracarida: Mysidacea) in relation to a variable environment

The euryhaline mysid Neomysis integer (Peracarida: Mysidacea) is a common member of the hyperbenthos of the upper reaches of European estuaries. In the East Looe River Estuary (Cornwall, England), this species experiences extensive tidal and seasonal changes in temperature (3 to 15 °C) and salinity (1 to 34‰). In this investigation, the effects of temperature (5, 10 and 15 °C) and salinity (1, 10, 20 and 30‰) on the oxygen consumption of male and female N. integer are reported, and are related to field measurements to identify the adaptive responses of the respiratory physiology to such a variable environment. The general responses were similar for each sex; however, at any given temperature/salinity combination, male N. integer consumed more oxygen than females. The general trends were increased oxygen consumption with increasing temperature (Q₁₀ values ranged from ∼1.7 to 2.5) and decreased oxygen consumption with increasing salinity. Temperature and salinity interacted at high water antagonistically to minimise changes in mysid oxygen-consumption. When related to tidal fluctuations in temperature and salinity experienced by N. integer inhabiting the East Looe River Estuary, the results reveal how the respiratory physiology of this species is adapted to its variable environment. Received: 16 June 1998 / Accepted: 15 December 1998     

Feeding in a larval fish assemblage: the nutritional significance of an estuarine plume front

Gut fullness of larval fishes was used to determine the nutritional significance of an estuarine plume front off Botany Bay, SE Australia, on three days in March/April 1990. Fishes were captured in three different water masses (estuarine plume, front and shelf water), each separated by 200 m, using a 260 m mesh purse-seine net. Overall, the gut-fullness index (GFI) of 260 fish larvae combined from eight families (Gerreidae, Mugilidae, Mullidae, Sparidae, Blenniidae, Kyphosidae, Monodactylidae, Pomacentridae), was significantly greater in the plume and front water on two of the sampling occasions, but no difference was detected on the third occasion. Trends in GFI among families were inconsistent with respect to the front. The mugilid (Liza argentea) fed equally and abundantly in all water masses (gut fullness>90%), while the kyphosid (Kyphosus spp.) had a significantly greater GFI in the plume compared to the shelf water. In general, the response of GFI to the front varied between dates and amongst taxa. Diet analysis showed that mugilids selected for copepod nauplii within the plume and shelf water [alpha selectivity index (ASI) of (0.22 to 0.98)] and for harpacticoid copepods within the front (ASI=0.92), representing characteristic diets identified by canonical discriminant analysis. Kyphosids selected positively for copepod nauplii in all three water masses on both occasions (ASI>0.96). In general, diets of larval fish were taxon-specific, and responded variably to the three habitats. Comparison of diets across water masses was complicated by the large number of families, which rarely occurred in all water masses. The nutritional significance of an estuarine plume front varies between species of larval fish, and there are no obvious trends that can be applied to the larval fish community in general.     

DDT isomers and metabolites in the environment: an overview

It is known for decades that the isomeric composition of organic pollutants can be influenced substantially by environmental processes such as biotransformation or transfer between compartments. This accounts also for the pesticide 2,2,-bis(4-chlorophenyl)-1,1,1-trichloroethane, better known as p,p′-DDT, and its accompanied substitution isomer 2-(2-chlorophenyl)-2-(4-chlorophenyl)-1,1,1-trichloroethane (o,p′-DDT). Although many studies followed the environmental fate of DDT, only very few publications reported on quantitative data of both o,p′- and p,p′-isomers. Therefore this condensed review describes evidence for remarkable changes and shifts in o,p′-/p,p′-ratios of DDT-related compounds. The application of isomer-specific analysis remains dominantly on emission source apportionment, for example, to differentiate DDT and dicofol emission. Only very few studies linked observed isomer shifts to aspects of environmental processes, such as (1) volatility from soil to air, (2) environmental stability in soil or (3) bioaccumulation in fishes. Additionally, several studies failed to use isomer-specific interpretation in order to obtain more detailed insight into environmental processes, for example, for observed isomer shifts during air–water fluxes. The o,p′-/p,p′-ratios of DDT and its main metabolite DDD have been detected more or less on the same level, whereas the isomers of the second main metabolite DDE were definitely depleted by the o,p′-isomer in all environmental compartments, indicating a general isomer-specific differentiation during DDT metabolism.     

Respiratory metabolism of crabs from marine and estuarine habitats: an interspecific comparison

The gill-surface area and R-T (metabolic rate-temperature) response during aquatic and aerial respiration of three different species of crabs, the subtidal Scylla serrata, the intertidal Sesarma quadratum, and the supratidal Ocypoda platytarsis, were compared. Scylla serrata has the highest, Sesarma quadratum the second highest, and O.platytarsis the lowest gill-surface area. Oxygen-consumption values in water under different ambient partial pressures of oxygen indicate that all three species display equally efficient aquatic respiration despite variation in gill area and number. There is no coherent R-T response either inter- or intra-specifically. However, the ability to extract oxygen from the surrounding medium (respiratory efficiency) conditions the R-T response in the different species. The R-T trends in aquatic respiration of Scylla serrata and aerial respiration of O.platytarsis reveal a normal response to temperature. The proportion of aquaticoxygen uptake to total respiration in all sizes in both Scylla serrata and Sesarma quadratum remains more or less constant (between 88 and 92%), whereas in the supratidal O.platytarsis, this proportion decreases with increasing weight (from 88 to 64 %), indicating a progressive loss of gill respiration to total respiration. All three species are metabolically equally efficient; what one lacks in respiratory surface it compensates by respiratory efficiency.     

Allocation of nitrogen and carbon in an estuarine salt marsh in Portugal

Above and below-ground biomass and nitrogen and carbon composition ofSpartina maritima, Halimione portulacoides andArthrocnemum perenne, dominating species in plant communities of the lower, middle and higher salt marsh, respectively, were compared in an estuarine salt marsh in Portugal. Plant and soil nitrogen and carbon pools were estimated. For all three species root biomass was significantly higher (70–92% of total biomass) than above-ground biomass. The percentage of root biomass was related to the location of the plants in the marsh: higher values were found in plants growing in the lower salt marsh where the sediment was more unstable and subject to tidal action, which stresses the role of the roots as an anchor. For all three species nitrogen concentrations were highest in leaves, reflecting the photosynthetic role of the tissue. For carbon higher concentrations were found in the stems, with the exception ofS. maritima. In general, lower nitrogen concentrations were found in summer, which can be explained by dilution processes due to plant growth. For both nitrogen and carbon, higher concentrations were found in the soil surface layers. Higher soil nitrogen and carbon levels were associated with higher organic matter contents. Most of the nitrogen in the salt marsh occurred in the sediments (0–40 cm) and only ca. 5.7–13.3% of the total was found in the plants. The greater portion (76.5%–86%) of carbon was found in the sediment.     

Dietary contribution of the microphytobenthos to infaunal deposit feeders in an estuarine mudflat in Japan

The food sources of benthic deposit feeders were investigated at three stations in an estuarine mudflat (Idoura Lagoon, Sendai Bay, Japan) during July and August 2005, using δ¹³C and δ¹⁵N ratios. Sediment at the stations was characterized by low chlorophyll (chl) a content (0–1 cm depth, <4 μg cm⁻²) and the dominance of riverine–terrestrial materials (RTM) in the sediment organic matter (SOM) pool. Surface-deposit feeders (Macoma contabulata, Macrophthalmus japonicus, and Cyathura muromiensis) exhibited much higher δ¹³C values (−18.4 to −12.4‰) than did the SOM pool (<−25‰). A δ¹³C-based isotopic mixing model estimated that benthic diatoms comprised 45–100% (on average) of their assimilated diet, whereas RTM comprised a lesser fraction (29% maximum). The major diet of the deep-deposit feeding polychaetes Notomastus sp. and Heteromastus sp. was benthic diatoms and/or marine particulate organic matter (POM), with little RTM assimilated (39% maximum). The consumers appeared to lack specific digestive enzymes and to use detritus-derived carbon only after its transfer to the microbial biomass. The isotopic mixing model also showed that the dietary contribution of RTM increased slightly (15% maximum) in the vicinity of freshwater input, suggesting that spatial changes in RTM supply affect the dietary composition of deposit feeders. These results clearly demonstrate that deposit feeders selectively ingest and/or assimilate the more nutritious microalgal fractions in the SOM pool. Such adaptations may allow enhanced energy gain in estuarine mudflats that are rich in vascular plant detritus with low nutritive value.     

Relationship between phytoplankton cell size and the rate of orthophosphate uptake: in situ observations of an estuarine population

Orthophosphate uptake by a natural estuarine phytoplankton population was estimated using two methods: (1) ³²P uptake experiments in which filters of different pore sizes were used to separate plankton size-fractions; (2) ³³P autoradiography of phytoplankton cells. Results of the first method showed that plankton cells larger than 5 m were responsible for 2% of the total orthophosphate uptake rate. 98% of the total uptake rate occurred in plankton composed mostly of bacteria, which passed the 5 m screen and were retained by the 0.45 m pore-size filter. There was no orthophosphate absorption by particulates in a biologically inhibited control containing iodoacetic acid. Orthophosphate uptake rates of individual phytoplankton species were obtained using ³³P autoradiography. The sum of these individual rates was very close to the estimated rate of uptake by particulates larger than 5 m in the ³²P labelling experiment. Generally, smaller cells were found to have a faster uptake rate per m³ biomass than larger cells. Although the nannoplankton constituted only about 21% of the total algal biomass, the rate of phosphate uptake by the nannoplankton was 75% of the total phytoplankton uptake rate. Results of the plankton autoradiography showed that the phosphate uptake rate per unit biomass is a power function of the surface: volume ratio of a cell; the relationship is expressed by the equation Y=2x10^-11 X ^1.7, where Y is gP m^-3 h^-1 and X is the surface: volume ratio. These results lend support to the hypothesis that smaller cells have a competitive advantage by having faster nutrient uptake rates.     

Diurnal vertical migration of an estuarine cladoceran,Podon polyphemoides,in the Chesapeake Bay

Populations of the marine cladoceran Podon polyphemoides (Leuckart) in the Chesapeake Bay (USA) zooplankton typically occur in large patches, extending the length of the estuary. These patches are characterized by well defined maxima, which do not move seaward during a given season. During the day, the bulk of the population is situated in the upper part of the two-layered estuarine circulation, where it is subjected to transport by net non-tidal seaward current. One mechanism for maintaining the population within the estuary is provided by the diurnal behavior of the podonids, which migrate to deeper water during the night. The landward advection of the deeper currents, thus, helps to counterbalance seaward population loss during the day. The extent of the vertical movement of the population can be modified by various environmental factors, but a suggestive correlation with light was found. It is thought that the downward component of the migration is due to passive sinking.     

Population dynamics and secondary production in an estuarine population of Nephtys hombergii (Polychaeta: Nephtyidae)

From July 1978 to March 1980, a study was made on the distribution, population dynamics and secondary production of Nephtys hombergii Audouin et Edw. occurring in the sublittoral industrialised region of Southampton Water in south England. The distribution of the worm was related to the silt content and copper level of the sediment, the greatest densities of N. hombergii being found in sediment containing 60 to 100% silt. Breeding occurred at a low level throughout the year, with a maximum in July to September and November to January in the second year of growth. Spawning occurred when the oocytes measured 200m in diameter, and unshed gametes were resorbed. Annual production varied between 0.092 and 4.32 g C m^-2 yr^-1 (ash-free dry weight) and amounted to 1.9–39.4% of the total macrofaunal production at the sampling stations. The production:biomass (P:B) ratio of the species varied between 1.6 and 2.9.     

Population dynamics and secondary production in an estuarine population of Caulleriella caputesocis (Polychaeta: Cirratulidae)

Between July 1978 and March 1980, the distribution, population dynamics and secondary production of Caulleriella caputesocis St. Joseph, 1864, in Southampton Water, South England, were investigated. The distribution of the polychaete was related to amount of silt and copper-content of the sediment, the highest densities occurring in sediment containing 60 to 100% silt and less than 50 ppm copper. Breeding occurred at a low level throughout the year, usually when the worms were over one year old, with peak breeding between May and September. C. caputesocis in Southampton Water is monotelic, and oocytes measured 120 to 160 m in diameter at spawning. Unshed oocytes were resorbed. Annual secondary production varied between 0.01 and 5.90 g C m^-2 yr^-1 and P:B ratio of the species ranged between 0.84 and 7.09.     

Poecilogony in an estuarine opisthobranch: planktotrophy, lecithotrophy, and mixed clutches in a population of the ascoglossan Alderia modesta

A San Diego population of the opisthobranch mollusc Alderia modesta (Lovén, 1844) exhibits poecilogony, the presence of two development modes within a single species. In spring, half of the adults spawned masses containing ∼300 eggs with a mean diameter of 68 μm. After 3 d, these egg masses hatched planktotrophic veligers with a maximum shell dimension of 116 μm. The remaining adults spawned masses containing ∼30 eggs with a mean diameter of 105 μm. These egg masses hatched after 5 to 6 d, releasing lecithotrophic larvae with a maximum shell dimension of 186 μm. About 1% of field-collected adults produced mixed clutches containing a continuum of larval sizes, spanning the size extremes of planktotrophy and lecithotrophy and hatching larvae with a mean maximum shell dimension of 152 μm. Adults producing planktotrophic and lecithotrophic larvae were interfertile, and no hybrid breakdown was observed through the F3 generation. When starved, adults which previously produced only lecithotrophic larvae switched to producing planktotrophic larvae or mixed clutches with both planktotrophic and lecithotrophic larvae. Sequence-polymorphisms from a portion of the mitochondrial cytochrome c oxidase I gene support the conclusion that the two reproductive morphs represent a single species. Most of the lecithotrophic larvae and a small percentage of the larvae from mixed clutches were metamorphically competent within 3 d of hatching. A. modesta is the only molluscan species as yet known to have both planktotrophic and pelagic lecithotrophic development within a single natural population. Received: 14 August 1997 / Accepted: 11 April 1998     

Effects of physical disturbance on an estuarine intertidal community: field and mesocosm results compared

The response of an estuarine macrobenthic community to different intensities of physical disturbance was examined using comparable field and mesocosm experiments. In the field study, areas of sediment (2500 cm²) were disturbed by surface-raking at a range of disturbance intensities: no disturbance, once every 2 mo, once every month, once every 2 wk, once or three times every week. These disturbance regimes were also imposed on buckets of sediment (855 cm²) collected from the field study site and maintained within a mesocosm system. In the univariate data analysis, neither field nor mesocosm communities displayed any trends in relation to the frequency of disturbance. Diversity indices were generally reduced in the mesocosm communities compared with the field communities. Multivariate analysis indicated significant differences between the field communities subjected to the most intense disturbance and those communities disturbed every ±2 wk or less. These differences were attributed to reductions in the five numerically dominant species in the high- disturbance treatments. More statistically significant differences were observed between the different disturbance intensities in the mesocosm experiment than in the field study. The implications of differences between field and mesocosm experiments results are discussed. Received: 14 June 1999 / Accepted: 3 December 1999     

Primary productivity and particulate organic matter flow in an estuarine mangrove-wetland in Hong Kong

Net primary productivity and organic matter flow of a mangrove-dominated wetland was estimated by following production and detritus dynamics in a tidal pond in north west Hong Kong in 1986–1988 (9.1 ha). Total productivity was 12.47 t dry wt ha^–1 yr^–1, of which >90% was from emergent macrophytes (the mangroveKandelia candel and the reedPhragmites communis). High turbidity and high summer temperatures probably limited respective production by phytoplankton and benthic macroalgae (dominated byEnteromorpha crinata). Despite the high total productivity, little detritus was exported from the emergent macrophyte stands, due to the low inundation frequency. This created a net water column carbon deficit which was provided for by the high organic matter import (mean = 4.42 g ash free dry wt m^–2 d^–1) from the incoming water. This same sediment and particulate organic carbon input giving a high accretion rate of 1.7 cm yr^–1 was probably also the force behind progressive dis-coupling of emergent macrophyte production from water column consumers. This resulted in a tendency to retain production in the emergent macrophyte stands while the water column community increasingly relied on allochthonous carbon. This shift from a net exporter to a net importer of carbon in landward wetlands is probably characteristic of the transition into nutrient-conservative terrestrial systems.     

Lead in New Orleans soils: New images of an urban environment

This paper describes a survey of lead in soil and computer generated maps that have been derived for New Orleans, Louisiana. The soil survey included streetside, houseside and open space samples. Because the survey covered every census tract in the metropolitan area it was possible to construct a computer-generated map of the distribution of lead dust in the soils of the urban environment. The data base consists of coordinates, site characteristics and lead analytical results of 3,704 soil samples. The resulting graphics show peaks of lead ranging from 600–1,200 g per g in the streetside soil of the inner-city and a steeply declining slope to the suburban areas of the city where the lead content of streetside soils is less than 75 g/g. In the inner-city, the amount of lead in soils found near building foundations is 10 to 20 times higher than the soils adjacent to streets where the median lead content of soils is over 300 g g^–1. In areas surrounding the city core (mid-city), the amount of lead next to the foundation and adjacent to the street are equivalent with medians of 110 g g^–1. In suburban locations, the median lead content of soil along streetsides is 86 g g^–1. Soils adjacent to surburban foundations has a median Pb content of 50 g g^–1. The lowest median lead content in soil is found in open spaces, ranging from 212 to 40 to 28 g g^–1, respectively, for the inner-city, mid-city, and suburbs. These observations are consistent with the production and consumer use of lead-based paint and leaded-fuels within the modern city.     

Huddling up in a dry environment: the physiological benefits of aggregation in an intertidal gastropod

In many intertidal gastropods, the formation of aggregations and closing of the opercular opening are behaviors commonly assumed to be associated with water conservation and maintenance of body temperature during tidal emersion periods. In the laboratory, we quantified the relationship between these two behaviors in a littorinid snail common to the north-central shores of Chile, Echinolittorina peruviana, and evaluated any benefit of these behaviors during desiccating conditions. We predicted that solitary individuals would maintain their opercula open for less time than aggregated snails when exposed to drier conditions due, at least in part, to differences in evaporative water loss. In laboratory trials, where relative humidity was manipulated, we observed that aggregated snails maintained their opercula open for longer periods of time than solitary snails under increasingly drier conditions. These results, together with observations of body temperature, suggest that aggregated animals may able to maintain gaseous exchange with their environment for longer periods of time than solitary individuals in response to desiccation stress. Our results suggest an interactive effect of two behaviors that confer physiological benefits when confronted with extreme physical conditions experienced during periods of emersion.     

Environmental risk assessment of cobalt and manganese from industrial sources in an estuarine system

A total of 74 samples of soil, sediment, industrial sludge, and surface water were collected in a Mediterranean estuarine system in order to assess the potential ecological impact of elevated concentrations of Co and Mn associated with a Terephthalic (PTA) and Isophthalic (PIPA) acids production plant. Samples were analyzed for elemental composition (37 elements), pH, redox potential, organic carbon, and CaCO₃ content, and a group of 16 selected samples were additionally subjected to a Tessier sequential extraction. Co and Mn soil concentrations were significantly higher inside the industrial facility and around its perimeter than in background samples, and maximum dissolved Co and Mn concentrations were found in a creek near the plant’s discharge point, reaching values 17,700 and 156 times higher than their respective background concentrations. The ecological risk was evaluated as a function of Co and Mn fractionation and bioavailability which were controlled by the environmental conditions generated by the advance of seawater into the estuarine system during high tide. Co appeared to precipitate near the river mouth due to the pH increase produced by the influence of seawater intrusion, reaching hazardous concentrations in sediments. In terms of their bioavailability and the corresponding risk assessment code, both Co and Mn present sediment concentrations that result in medium to high ecological risk whereas water concentrations of both elements reach values that more than double their corresponding Secondary Acute Values.     

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