Role of herbivory in controlling phytoplankton abundance: annual pigment budget for a temperate marine fjord

An annual pigment budget was constructed for Dabob Bay, Washington (USA) by comparing the downward vertical loss of phytoplankton pigments (chlorophyll and pheopigments) to the production of chlorophyll within the euphotic zone. The vertical flux of pigments was measured with sediment traps deployed at intervals of 1 to 6 wk over a 2.5 yr period yielding 763 d of trap exposure (28 November 1978–16 June 1981). The production rate of chlorophyll was calculated from measurements of algal specific growth rates, chlorophyll (chl) crops, primary production (as carbon) and appropriate C: chl ratios. Sixty one to 77% of the annual chlorophyll production was accounted for by the vertical flux of pheopigments resulting from herbivorous zooplankton grazing (macrozooplankton). Algal sinking, represented by downward chlorophyll flux, accounted for only 5 to 6% of the annual chlorophyll production. The remaining fraction of chlorophyll production was estimated to be consumed by small herbivores (microzooplankton), whose fecal material contributes to the suspended pool of pheopigments found in the euphotic zone. The suspended pheopigments are continuously removed by photodegradation. In Dabob Bay, the major process controlling phytoplankton abundance is zooplankton grazing and it appears that the ultimate fate of most phytoplankton is to be consumed by herbivores.     

Variability and its effect on the 24h chlorophyll budget of a small marine basin

In an intensive study (lasting 25 h) of the production, export and grazing of phytoplankton in a small marine basin, it was found that 58% of the production (11% of the total standing stock) was lost by exchange with the sea and 34% was consumed by grazing of zooplankton. The measured production of phytoplankton could be balanced, to within a few percent, against grazing, export, and a small, measured, net change in the total standing stock of the basin. Large variations were observed in concentrations of chlorophyll and zooplankton at the mouth of the basin over the 25 h period. These variations were associated with changes in the height of the tide, but were about 2^1/2 h out of phase with it. Strong negative correlations were observed between chlorophyll and transport, such that only 35% of the chlorophyll exported was exchanged via the mean flow, while 65% was exchanged via the fluctuations. The correlation was even more striking with zooplankton, for which virtually all the export was associated with the fluctuations in the transport. Time series observations in the centre of the basin revealed considerable short-term variability in both chlorophyll and zooplankton, but the variations were smaller than those observed at the mouth of the basin, and the phase lag with the tide was longer. The variability studies enable suggestions to be made about more economical design of sampling programs, but illustrate the difficulty of providing verification data for any continuous model of primary production in such a basin.Bedford Institute Contribution No. 231.Canadian Contribution to IBP No. 97.     

Vertical distribution of zooplankton glutamate dehydrogenase in relation to chlorophyll in the vicinity of the Nantucket Shoals

The vertical distributions of chlorophyll, and zooplankton glutamate dehydrogenase (GDH) and respiratory electron transport system (ETS) activities were measured at five stations in the vicinity of the Nantucket Shoals using a submersible pump-profiling system. Distinctly lower zooplankton GDH activities and chlorophyll concentrations were found at a station showing little vertical stratification as compared to the other stations where vertical stratification was more pronounced. Maximum GDH activity was observed at or immediately above the chlorophyll maximum at the stratified stations. Respiration to excretion (O:N) ratios determined from the enzyme data were lowest in or just above the chlorphyll maximum, suggesting proteolytic catabolism at those depths. Both features had previously been observed in the oligotrophic Gulf of Mexico. Ammonium excretion determined from GDH activity was estimated to provide 20 to 40% of the total algal nitrogen requirement.Bigelow Laboratory Contribution No. 82016     

Inter- and intra-specific diurnal habitat selection of zooplankton during the spring bloom observed by Video Plankton Recorder

Diel vertical migration (DVM) is a common behavior adopted by zooplankton species. DVM is a prominent adaptation for avoiding visual predation during daylight hours and still being able to feed on surface phytoplankton blooms during night. Here, we report on a DVM study using a Video Plankton Recorder (VPR), a tool that allows mapping of vertical zooplankton distributions with a far greater spatial resolution than conventional zooplankton nets. The study took place over a full day–night cycle in Disko Bay, Greenland, during the peak of the phytoplankton spring bloom. The sampling revealed a large abundance of copepods performing DVM (up during night and down during day). Migration behavior was expressed differently among the abundant groups with either a strong DVM (euphausiids), an absence of DVM (i.e., permanently deep; ostracods) or a marked DVM, driven by strong surface avoidance during the day and more variable depth preferences at night (Calanus spp.). The precise individual depth position provided by the VPR allowed us to conclude that the escape from surface waters during daytime reduces feeding opportunities but also lowers the risk of predation (by reducing the light exposure) and thereby is likely to influence both state (hunger, weight and stage) and survival. The results suggest that the copepods select day and night time habitats with similar light levels (~10^?9 μmol photon s^?1 m^?2). Furthermore, Calanus spp. displayed state-dependent behavior, with DVM most apparent for smaller individuals, and a deeper residence depth for the larger individuals.     

Post-bloom grazing byCalanus glacialis,C. finmarchicus andC. hyperboreus in the region of the Polar Front,Barents Sea

The plankton community in the Polar Front area of the Barents Sea was investigated during a cruise from 14 to 28 July 1987. The colonial algaePhaeocystis pouchetii andDinobryon pellucidum dominated the phytoplankton. Depth integrated carbon assimilation rates varied from 190 to 810 mg C m^–2 d^–1. A high carbon:chlorophyll ratio (which varied from 123 to 352) prevailed at the three stations investigated, which may relate to facultative heterotrophic behaviour byD. pellucidum. The herbivorous zooplankton community was dominated byCalanus glacialis, C. finmarchicus, andC. hyperboreus. Maximum zooplankton biomass was found in the same depth strata as phytoplankton chlorophyll maximum. The herbivorous copepod populations did not display consistent day-night vertical migration patterns. Phytoplankton consumption rates of the various life stages were estimated from the turnover rate of plant pigments in the gut. The gut defecation rate constant (R) varied from 0.014 to 0.027 min^–1 at 0°C in copepodites (Stage II to adult female) ofC. glacialis, independent of developmental stage.Calanus spp. community carbon ingestion rates calculated from particulate carbon:chlorophyll ratios, were 10, 65 and 400% of daily phytoplankton carbon fixation rates at Stations 1, 2 and 3, respectively.     

海水混合和层化对叶绿素a垂直分布的影响

根据长江口海区、近岸浅水区、黄海冷水团海区和水深超过200m的陆架区等四个区域的定点调查获得的数据，分析了温度、盐度与叶绿素a垂直分布的相互关系。结果表明，长江口海区，陆源径流与海水混合不充分，表层营养盐含量较高，表层叶绿素a含量高于中下水层。近岸浅水区的苏北近岸海水垂向没有温跃层和盐跃层，叶绿素a的垂向分布也均匀，东海西部沿岸出现逆温跃层和逆盐跃层，海水垂直混合不充分，叶绿素a含量(6．72mg／m^3)在10m深水域最高。黄海冷水团海区海水的垂直混合不充分，叶绿素a的垂直变化显著，高值区出现在温跃层下方(4．37mg／m^3)。水深超过200m的陆架区，温度的阶梯状结构、营养盐跃层、光照等因素共同导致整个水层叶绿素a含量普遍较低。同时，结合历史资料分析认为，海水的混合、温度和盐度的层化将影响营养盐的浓度和分布，从而影响海水中叶绿素a的垂直分布。     

Influence des conditions hivernales sur les productions phyto-et zooplanctoniques en Méditerranée Nord-Occidentale. V. Biomasse et production zooplanctonique — relations phyto-zooplancton

The cruise “Mediprod I” of the R.V. “Jean Charcot” covered an area of the Mediterranean Sea situated North of the 40th parallel and West of Corsica; during two 15-day legs, the first when Winter conditions ended, in March, the second in Spring conditions, in April, a 48-station network was surveyed as to primary and secondary production, as well as hydrological conditions. The first survey revealed a higher concentration of zooplankton in hydrologically stable areas, especially in the South-East, although zooplankton values were rather low throughout the whole area surveyed. The total zooplankton per unit surface was lower at the edges of the surveyed area, where phytoplankton was scarce. An increase in zoo-plankton biomass was observed between the two surveys, mostly in the central areas and near the surface. An important difference between both legs is in the proportion of organisms of different sizes collected by the Clarke-Bumpus sampler: the-200 μ:+200 μ organisms ratio, which is around 1 during the first leg, is much lower during the second leg. Two methods were used in estimating the biomass: the +200 μ fraction of a Clarke-Bumpus sample (Cl-B towed with a 50-μ net fitted on the sampler) was collected by sieving the sample through a 200-μ mesh nylon; standard vertical WP 2 hauls were performed (200 μ mesh). Both roughly show the same zooplankton (weight per unit surface) distribution pattern. However, higher estimations of the total biomass of the volume investigated were sometimes provided by the Clarke-Bumpus method during the first leg, probably due to the distribution of the animals according to their size-classes. A graph of chlorophyll versus zooplankton for surface waters suggests that zooplankton has a limiting effect on the development of phytoplankton in April only. Apparent growth rates of cooplankton are less than those for populations in the laboratory or enclosed environments. Values of the mean secondary production vary from 18 mg C.m^-2.day^-1 for the first leg, to 230 mg C.m^-2.day^-1 for the second leg. Estimations of net efficiency for energetic transfer between phyto- and zooplankton lie between 7 and 26%. As far as our hypotheses as regards physiological coefficients are valid, we can assume that the effect of grazing relative to primary production is greater in the border areas than in the central area, thus increasing the contrast between both areas with time. Phosphorus excretion rate by zooplankton seems to be less than that measured in the Atlantic Ocean. We suggest that in Spring zooplankton excretion is not the main phosphorus recycling process. Primary productivity measurements, apparent growth coefficients, and estimated grazing rate have been used to calculate the expected mean biomass per unit area during the second leg. A 7% loss of phyto- and zooplankton from the upper 100-m layer must be assumed to explain the observed biomass variation. Vertical mixing and sinking of surface water on an isopycnal slope are responsible for such loss, which can also affect, to an undetermined degree, the phosphorus stock introduced into the surface layer by the Mediterranean deep-water formation mechanism. We suggest that nearly half the total loss of phytoplankton from March to April is attributable to animal grazing.     

Differences in productivities between the Great Australian Bight and the Gulf of Carpentaria,Australia, in summer

Phytoplankton standing crop (chlorophyll a) and primary productivity were recorded, and zooplankton biomass was estimated in the two large bays of Australia, the Great Australian Bight on the south coast (December, 1965) and the Gulf of Carpentaria on the north coast (December, 1968). In the Gulf of Carpentaria, the phytoplankton standing crop (average, 27.3 mg chlorophyll a m^-2) and primary productivity (average, 133.1 mg C m^-2 h^-1), as well as zooplankton biomass (average, 305.3 mg wet weight m^-3) are much higher than in the Great Australian Bight (12.1 mg chlorophyll a m^-2, 18.2 mg C m^-2 h^-1, 7.1 mg wet weight m^-3, respectively). The unexpectedly low productivity values in the Great Australian Bight are attributable to environmental conditions of this bay, which obtains neither replenishment of nutrients from the land nor receives upwelling of deep water.     

Observations of copepod feeding and vertical distribution under natural turbulent conditions in the North Sea

We present results of simultaneous measurements of turbulent-dissipation rate, zooplankton vertical distribution and copepod gut pigments in the northern North Sea. Analysis shows that some, but not all, copepods (by species, sex and stage) exhibit significant dependence on turbulence in respect to vertical distribution and feeding rate. Oithona similis (female and copepodite stages) exhibits an avoidance of the surface layer when turbulence is strong there. For the range of turbulence (10⁻⁷ to 10⁻³m² s⁻³) and ambient chlorophyll concentration (0.5–0.8 μg l⁻¹) encountered, Calanus spp. and Metridia lucens exhibited a significant negative response in feeding-rate index with increasing turbulence. Centropages typicus and Pseudocalanus spp. also exhibited a negative response but of less significance. Received: 12 October 2000 / Accepted: 11 December 2000     

Nutrient regeneration by zooplankton during a red tide off Peru,with notes on biomass and species composition of zooplankton

During March and April 1976, a red tide, dominated by the dinoflagellate Gymnodinium splendens Lebour, developed in the vicinity of 15°06'S and 75°31'W off Peru. At the height of the bloom, the euphotic zone was 6 m deep and the chlorophyll a at the surface was 48 g l^-1. A daily collection of zooplankton at 09.00 hrs showed large fluctuations of biomass, from 0.2 to 3.84 g dry weight m^-2 in a water column of 120m. Copepodids and nauplii dominated the collections. During a period of reduced wind, the adult copepods were a mixture of the species characteristic of the coastal upwelling system and the neritic species associated with more northerly, tropical waters. Nitrogen regeneration by the zooplankton varied with the development of the bloom, the type of zooplankton dominating the experiment, and biomass fluctuations, but never accounted for more than 25% of the nitrogen uptake by phytoplankton.     

Distribution of lipopolysaccharide,an indicator of bacterial biomass,in subtropical areas of the sea

Lipopolysaccharide (LPS) concentrations, an indicator of bacterial biomass, were determined in the South China Sea and the Pacific Ocean. The distribution patterns of LPS were compared with those of chlorophyll a (Chl a), zooplankton biomass and the concentrations of several nutrients. LPS and total bacterial numbers in seawater were correlated with each other with a correlation coefficient (r) of 0.84 at Stations 7, 8 and 10. Diurnal fluctuation of LPS was negligible, but Chl a varied slightly in the vertical water column. Zooplankton stayed at a depth of around 400 m during the daytime and ascended quickly to the surface (0–50 m) early in the evening. The profiles of LPS and Chl a were negatively correlated to each other in the water layers above the Chl a maximum peak (r=-0.74; excluding the samples from 75 m at Station 7 and 10 m at Station 11 due to inadequate data for the statistical analysis). LPS and zooplankton biomass during the night-time, in contrast, paralleled each other at 5 stations surveyed (r=0.71). The presence of zooplankton resulted in an increase in bacterial numbers in the seawater in vitro. Based on these results, the factors controlling the occurrence and abundance of bacteria and phyto- and zooplankton in the pelagic sea are discussed.     

Effects of buoyancy, transparency and zooplankton feeding on surface maxima and deep maxima: Comprehensive mathematical model for vertical distribution in cyanobacterial biomass

Inhomogeneous vertical distributions of the cyanobacterial biomass are widely observed during the summer season in stratified lake ecosystems. Among these are surface maxima characterized by surface scum formation and deep or subsurface maxima also known as deep chlorophyll maxima (DCM). The former occurs at the epilimnion in eutrophic lakes, and are usually caused by colonial cyanobacteria such as Microcystis. On the other hand, the latter occurs at the metalimnion and the upper part of the hypolimnion near the thermocline in oligotrophic lakes, and are referred to filamentous cyanobacteria such as Oscillatoria. The aim of this paper is to present a simple mathematical model that can simultaneously describe these phenomena including the annual and diurnal variations, emphasizing the roles of buoyancy regulation, transparency of the lake and zooplankton feeding on cyanobacteria. According to our computer analyses, the increased buoyancy, the low clarity of the lake and the low rate of zooplankton feeding take significant roles in formation of surface maxima, while the reversal of these factors makes deep maxima predominant. Our two-component model with nutrients and cyanobacteria can distinguish between two phenomena by changing the parameters for these factors, without altering the model itself.     

Relationship between the zooplankton,phytoplankton, particulate matter and dissolved free amino acids in the Celtic Sea

Estimates of the biomass of zooplankton, phytoplankton and particulate matter collected in the Celtic Sea during mixed-water conditions (on 8 and 9 April 1983) were compared to the concentration and diversity of sixteen dissolved free amino acids (DFAA) measured in seawater and in particles. During a day profile, variations of dissolved amino acids with depth reflected the feeding activity of copepods. The relationship was not apparent in a night profile and other processes, such as heterotrophic utilization of dissolved nitrogen by microorganisms, were thought to be involved. The ratios of total DFAA concentration (nM litre^-1) in the particulate phase over the concentration in seawater ranged from 1 to 200 within the water column. Of the sixteen amino acids measured, ornithine, a decomposition product of arginine, was responsible for more than 70% of the total concentration of DFAA in seawater. In the particles, phenylalanine ranged from 30 to 88% of total DFAA. In seawater this amino acid occurred in the 20 to 40 m depths (1.3 to 9.9% of total DFAA) in the day profile and at 5 m (12.4%) and 80 m (6.4%) in the night profile. Previously it has only been found in very low concentrations (<5%) in seawater, and its presence is considered to be the result of zooplankton feeding.     

The energetics and evolution of intraspecific predation (egg cannibalism) in the anchovy Engraulis capensis

Anchovy feeding activity (based on samples collected around the South African coast between 1986 and 1988) seems to peak twice daily, with the major peak between 17:00 and 19:00 hrs. Mean daily consumption in the field was estimated at 3.45% of body wt d^-1. Since the various food types available to anchovy differ in their caloric values, this figure might be expected to vary according to the type of diet available to the fish. To obtain a diet energetically equivalent to that measured in the field anchovy feeding exclusively on phytoplankton would need to consume the equivalent of 5.8% of their body wt d^-1 and those feeding on zooplankton 3.5 to 4.5% of the body wt, while fish feeding on their own eggs would require only 2.5% of their body wt daily. Feeding on eggs at peak spawning on the spawning grounds —where there is limited alternative food (i.e., zooplankton) available — seems adequate to support reproduction. In selection at the population level, cannibalistic fish benefit from homeostatic self-regulation.     

Zooplankton biomass and indices of grazing and metabolism during a late winter bloom in subtropical waters

The development of the so-called late winter bloom in subtropical water was studied in an oceanic area north of the Canary Islands from January to May 2000. Zooplankton was sampled at short-term intervals (1–4 days) during the bloom (January–March), and biomass, indices of grazing (gut fluorescence) and metabolism (electron transfer system activity, ETS) were measured in four different size fractions (100–200, 200–500, 500–1000 and >1000 µm). During the bloom, ETS activity and gut fluorescence increased before the development of zooplankton biomass. At the end of February, the presence of an impressive cloud of dust formed in the Sahara desert was related to an increase in chlorophyll and small zooplankton a week later. The increments in biomass were the consequence of consumption by zooplankton as inferred from the indices of grazing and metabolism. Estimated grazing from gut fluorescence and gut evacuation rates during the period of study accounted for 55% of the assessed total ingestion from respiration and normal values of assimilation, showing the importance of the non-pigmented food in the diet of zooplankton in these waters. In contrast, the sharp decreases in zooplankton biomass observed during the bloom appeared during the dark period of the moon, the days in which the diel vertical migrants reach the shallower layers, in agreement with previous works in the area. Thus, the development of the late winter bloom in this region is suggested to be driven by the interplay between resource and consumer controls.Communicated by S.A. Poulet, Roscoff     

Diel vertical migration bySalpa aspera and its potential for large-scale particulate organic matter transport to the deep-sea

In mid-summer 1975 throughout the Western Slope Water of the North Atlantic Ocean, massive numbers ofSalpa aspera performed a diel vertical migration of at least 800 m. This resulted in a movement of 85 to 90% of the total zooplankton biomass out of the upper 500 m during the day. Fecal pellet production and losses from this salp population were estimated to contribute approximately 12 mg C m^-2 day^-1 to the deep planktonic and benthic populations. If all this organic matter reached the deep-sea floor, it would represent over 100% of the daily deep-sea benthic infauna energy requirements.     

Description and use of an improved method for determining estuarine zooplankton grazing rates on phytoplankton

A method of rapidly determining zooplankton grazing rates on natural mixed phytoplankton populations using ¹⁴C is described. The method simplifies the design of grazing experiments as the grazing time can be kept short enough to prevent recycling of the isotope, and growth of the phytoplankton substrate. Very high specific activity, ¹⁴C-labelled phytoplankton concentrated either by centrifugation or sieving, may be used either as the sole grazing substrate, or as a tracer in natural mixed phytoplankton. Zooplankton, confined in glass jars at either ambient, or higher than ambient concentrations, are permitted to feed on the phytoplankton for periods of 30 min and 2 h, and are then separated by sieving. The zooplankton community grazing rate, or, if the samples are sorted into species, the individual species grazing rates, can be determined after scintillation counting of the zooplankton. The rate of appearance of ¹⁴C-labelled phytoplankton in the zooplankton is an estimate of the grazing rate, and the slope of the line joining the grazing rates at various phytoplankton concentrations gives an estimate of the grazing rate constant for the zooplankton population. The method provides a quick way of obtaining both zooplankton population, and individual species grazing rates on natural mixed phytoplankton. In two experiments, labelled phytoplankton was used as the sole grazing substrate in concentrations ranging between 0.4 and 5 times ambient levels. Grazing rate constants, for net-caught zooplankton concentrated to 46 times (Experiment 1) and 28 times (Experiment, 2) ambient estuarine levels were-0.14and-0.12 of the phytoplankton standing stock per day, respectively. There was a linear increase in the amount of phytoplankton grazed with an increase in phytoplankton concentration up to four times ambient phytoplankton levels. When tracer amounts of labelled phytoplankton were added to samples containing both phytoplankton and zooplankton at ambient concentrations the grazing rate constants were-0.28 and-0.42 of the phytoplankton standing stock per day. We conclude that zooplankton grazing was the major control factor of phytoplankton population size during October–November 1975 in South West Arm, Port Hacking, near Sydney, Australia.     

Behavioral responses of larvae of the crab Rhithropanopeus harrisii (Brachyura: Xanthidae) during diel vertical migration

Larval Rhithropanopeus harrisii (Gould) show nocturnal vertical migration. Larval behavioral responses to different rates of increase and decrease in light intensity were measured in an apparatus with a natural angular light distribution. A central objective was to establish whether phototaxis actually participates in vertical migration. At sunset the level of light adaptation controlled the readiness of the larvae to migrate, while an ascent was initiated by a preductable relative decrease in intensity (e.g. 4.0x10^-3s^-1). Rates of relative decrease around sunset would evoke continuous upward swimming. Gravity was the orienting cue and there was no change in swimming speed during the ascent. At sunrise, the larval descent was initiated by exposure to an absolute light intensity of about 0.23 log unit above the lower visual threshold. Light served as an orienting cue, as larvae descended by a negative phototaxis. Thus, phototaxis is not a laboratory artifact and does participate in vertical migration. A consideration of behavioral responses of other crustacean zooplankton indicates there is considerable variation in the initiating and directing cues for vertical movements. The variety of behavioral responses of R. harrisii suggests that a synthesis of hypotheses about migration may provide the proper basis for explaining the mechanisms underlying diel vertical migration.     

A method to determine in situ zooplankton grazing rates on natural particle assemblages

In situ zooplankton grazing rates on natural particle assemblages were stimated by measuring zooplankton uptake of labelled autotrophic (with Na¹⁴CO₃) and heterotrophic (with [methyl-³H]-thymidine) particulate matter in 1-h incubations in clear, Plexiglas, Haney chambers. The in situ grazing rates are in the same range as those measured for zooplankton in the laboratory using standard particle counting techniques. A negative selection coefficient for ³H-labelled particles indicated a lower filtration efficiency or avoidance of these particles by zooplankton.     

A plankton study in the eastern Mediterranean Sea

A synoptic study of the phytoplankton and zooplankton distribution in the eastern Mediterranean Sea was carried out in the summer of 1965 during a cruise of R.V. “Pillsbury”. The phytoplankton maximum was normally found at 100 m, below the 1% level of surface light, with a frequent smaller peak at 40 m. In shallower waters, the main peak was at 80 m. The greatest number of phytophagous crustaceans was recorded at, or close to, the phytoplankton maximum. Depth relationships of phytoplankton and functionally phytoplanktonic species are discussed. Zooplankton was collected from depths down to 4,400 m, the deepest cast being in the Rhodes Deep. The most common species and genera of some selected groups of zooplankton were identified and their distribution considered in the light of some previous cruises in part of the area. The presence of Acantharia containing zooxanthellae with chlorophyll at, and below, 4,000 m was recorded for the first time. Potentially photosynthetic Ceratium vultur and C. carriense (Dinophyceae), both species in active division, were also found at these depths, as well as Halosphaera viridis (Prasinophyceae), which was recorded down to a depth of 1,000 m.