The heterotrophic phase of plankton succession in the Japan Sea

The vertical structure, composition and productivity of a plankton community was studied in the Japan Sea in June, 1972 during a period of thermocline formation; the parameters measured were: phytoplankton production and biomass; number, biomass, and production of planktonic bacteria; biomass of phagotrophic flagellates, ciliates and remaining microzooplankton. The concentration of micro- and mesozooplankton attained a basic maximum in a layer near the upper part of the thermocline. The biomass and calculated production of the heterotrophic part of the community exceeded considerably the amount of primary production. The heterotrophic phase of the seasonal succession of a plankton community in a temperate sea is described, when heterotrophic metabolism and production predominate. Heterotrophs at this stage use mostly energy from organic matter accumulated during the previous spring phytoplankton bloom.     

Continuous plankton records: The occurrence of apostome ciliates (Protozoa) on Euphausiacea in the North Atlantic Ocean and North Sea

The phoronts (resting cysts) of apostome ciliates have been recorded on Euphausia hemigibba, E. krohni, Meganyctiphanes norvegica, Nematoscelis megalops, Nyctiphanes couchi, Thysanoessa gregaria, T. inermis, T. longicaudata and T. raschi taken in the Continuous Plankton Recorder Survey of the North Atlantic Ocean and the North Sea. Variations in the percentage of host euphausiids infested by apostomes have been studied in relation to species, geography, body size and season. The percentage infestation was found to be related mainly to the body size of the hosts. The apostomes are believed to be species that develop either in the exuviae discarded exoskeletons or corpses of the euphausiids, or in the digestive systems of predators of the euphausiids.     

Changes in the phytoplankton community at Helgoland, North Sea: lessons from single spot time series analyses

In recent regime shift analyses, the phytoplankton compartment of the marine food web was essentially represented by phytoplankton color or chlorophyll concentration. A detection of changes directly at the species level is highly desirable. The Helgoland Roads data series, a collection of high frequency long-term time series comprising biological and physico-chemical components of the southern North Sea, allow such an investigation at the level of single species. Aiming at a detection and characterization of habitat and community changes in the observation period (1962 until the end of 2008), we selected six species as representatives of certain classes, for example, benthic or neritic species, and applied a combination of novel analysis methods—a fitness-based analysis of the realized niche, a bloom-triggered averaging and a Markovian analysis of co-occurrence and succession patterns—to related abundance time series and concurrent environmental parameter time series. We found a general trend toward enlargement of niche size and shifts of the niche position, interesting salinity patterns around bloom events of two species, and statistically highly significant changes of a phytoplankton community segment after 1965 and after 1998. Interpreting our observations in ecological terms leads to the formulation of testable hypotheses.     

The early summer bloom of dinoflagellates in the North Sea,with special reference to 1971

The south-western quarter of the North Sea (between the Wash and the River Tyne) has been investigated for the presence of dinoflagellates. Extensive net surveys were carried out in March, May and June 1971 and a number of other collections were made in the area. North of Flamborough Head, the spring bloom of diatoms was succeeded by a considerable growth of dinoflagellates, of which Dinophysis norvegica was the most abundant organism. Maximum counts of over 28 million dinoflagellates per m³ were recorded. Distribution charts are given for the most common species. A total of 61 species was found during the course of the survey.     

Density and composition of heterotrophic bacterial populations in North Sea sediments

The bacteria from 119 sediment samples collected off the Belgian and Dutch North Sea coast were enumerated on marine agar. Geographical and seasonal variations were observed. The colony formers were mainly obligate aerobes. Coliform organisms were relatively rare. An exhaustive analysis of a few samples revealed a great number of species belonging to different taxonomic groups. A survey of physiological and biochemical characteristics of 71 isolates is presented.     

Overwintering strategies of dominant calanoid copepods in the German Bight, southern North Sea

Abundance, stage composition and reproductive parameters (egg production, egg viability, proportion of spawning females) of the four copepod species Acartia clausi, Centropages hamatus, C. typicus and Temora longicornis were measured at the long term sampling station Helgoland Roads (German Bight, southern North Sea) from September 2003 to May 2004 to study their overwintering strategies. A. clausi was overwintering as females with arrested reproduction from November to January. T. longicornis, which is known to produce resting eggs in the North Sea, had a pelagic population with all developmental stages present during winter and reproductive rates closely related to food concentrations. Although their females produced eggs in response to ambient food conditions, both C. hamatus and C. typicus were rare in the pelagic. The C. hamatus population returned in May, probably from resting eggs, whereas C. typicus depended on advection. The Centropages species seemed to be less adapted to pelagic life in winter than A. clausi and T. longicornis. Sporadic occurrence of large numbers of nauplii and young copepodids of A. clausi and Centropages spp. pointed to different overwintering strategies or more successful survival in adjacent regions and advection of them into the waters around Helgoland island. While A. clausi was decoupled from environmental conditions in late autumn and winter, the other species were able to respond to variations in the food environment. Thus, egg production of T. longicornis increased during an unusual autumn diatom bloom.     

The role of environmental and spatial processes in structuring lake communities from bacteria to fish

We assessed the relative roles of local environmental conditions and dispersal on community structure in a landscape of lakes for the major trophic groups. We use taxonomic presence-absence and abundance data for bacteria, phytoplankton, zooplankton, and fish from 18 lakes in southern Quebec, Canada. The question of interest was whether communities composed of organisms with more limited dispersal abilities, because of size and life history (zooplankton and fish) would show a different effect of lake distribution than communities composed of good dispersers (bacteria and phytoplankton). We examine the variation in structure attributable to local environmental (i.e., lake chemical and physical variables) vs. dispersal predictors (i.e., overland and watercourse distances between lakes) using variation partitioning techniques. Overall, we show that less motile species (crustacean zooplankton and fish) are better predicted by spatial factors than by local environmental ones. Furthermore, we show that for zooplankton abundances, both overland and watercourse dispersal pathways are equally strong, though they may select for different components of the community, while for fish, only watercourses are relevant dispersal pathways. These results suggest that crustacean zooplankton and fish are more constrained by dispersal and therefore more likely to operate as a metacommunity than are bacteria and phytoplankton within this studied landscape.     

Egg production and feeding by copepods prior to the spring bloom of phytoplankton in Fram Strait,Greenland Sea

Reproductively activeCalanus hyperboreus (Krøyer) andC. glacialis Jaschnov were captured in the upper 100 m of Fram Strait (77° to 79°N) in late winter 1987. There was no evidence of a phytoplankton bloom; chlorophylla concentrations were uniformly low (<0.1 mg m^–3), and nitrate concentrations were uniformly high (>11.3 mg-at m^–3). Gut-fullness measurements indicated that females were ingesting very little. The maturation state of gonads of bothC. hyperboreus andC. glacialis indicated that 75% of females were in a ripe condition consistent with observed egg laying. The lipid content of females laying eggs was reduced in both species compared to that of females not laying eggs. InC. hyperboreus the reduction was 39% and inC. glacialis it was 44%. All the evidence suggests that bothC. hyperboreus andC. glacialis were laying eggs in late winter by using lipids stored previously; they were not relying on ambient concentrations of phytoplankton. The daily rate of egg laying byC. glacialis using lipids in late winter exceeded the rate reported for summer when ambient food supplies have been shown to be necessary. It is suggested that individuals, spawned well in advance of the spring bloom of phytoplankton, may comprise a major portion of the annual recruitment to the entire population ofC. glacialis in this area, and that their life cycle can be completed within 1 yr. NeitherMetridia longa (Lubbock) norC. finmarchicus (Gunnerus) laid eggs during this study.     

Production and respiration of overall plankton and ultraplankton communities at the entrance to the Gulf of Finland in the Baltic Sea

Primary productivity and respiration of the overall plankton community and of ultraplankton (organisms passing through a 3-m Nuclepore filter) were studied at the entrance to the Gulf of Finland during the growth season in 1982. Data of the respiration measurements from previous years are also presented. During the development of a diatom spring bloom, the algal component could be successfully separated from the bacterial component by size fractionation with a 3-m Nuclepore filter and thus the algal respiration could be approximated, being on the order of 10 to 20% of the gross production. After the phytoplankton spring maximum, bacteria played an important role in mediating the energy flow from phytoplankton exudates to higher trophic levels. Maximum values of 1 230 and 740 mg O₂ m^-2 d^-1 were recorded for overall and for ultraplankton respiration, respectively, during late July. High productivity values coupled with low phytoplankton biomass and low inorganic nutrient values were also recorded in late July, indicating effective nutrient regeneration and rapid turnover of the plankton community. During late summer, a considerable fraction (over 30%) of phytoplankton production was released as exudates, suggesting that much of the energy is channeled to higher trophic levels via bacterial pathways rather than by direct herbivorous grazing during this season. The summer development of phytoplankton community structure and functioning is strongly controlled by hydrographic conditions, i.e. by nutrient inputs via upwelling and by water temperature. A carbon budget for late summer indicated that bacteria may contribute only up to 50% of the overall respiration of the plankton community, which suggests that heterotrophs other than bacteria play an important role in nutrient regeneration. The present study stresses the importance of energy flow via the phytoplankton exudatebacteria-micrograzer pathway in relatively oligotrophic, brackish water ecosystems.     

Continuous Plankton Records: Seasonal cycles of phytoplankton and copepods in the North Atlantic ocean and the North Sea

Data from the Continuous Plankton Recorder survey of the North Atlantic Ocean and the North Sea are used to study geographical variations in the amplitude, duration and timing of the seasonal cycles of total phytoplankton and total copepods. It is shown that the distribution of overwintering stocks influences the distributions throughout the year. There is a relationship between the timing of the spring increase of phytoplankton and the amplitude of the seasonal variation in sea surface temperature. In the open ocean, the timing of the spring increase of phytoplankton corresponds with the spring warming of the surface waters. In the North Sea the spring increase occurs earlier, associated, perhaps, with transient periods of vertical stability, resulting in a relatively slower rate of increase. It is suggested that in the open ocean the higher rate of increase is under-exploited by copepods due to low overwintering stocks and longer generation times. Exceptionally early spring increases of phytoplankton off the west coast of Greenland and over the Norwegian shelf are probably associated with permanent haloclines. A high and late autumn peak of phytoplankton off the coast of Portugal may be associated with coastal upwelling.     

Water-soluble vitamins in natural plankton (copepods) during two consecutive spring blooms compared to vitamins in Artemia franciscana nauplii and metanauplii

Weight-specific contents of asorbic acid, thiamin, riboflavin, niacin, pantothenic acid, vitamin B₆, biotin, folate and vitamin B₁₂ were analysed in natural zooplankton collected from a sea water lagoon in a halibut fry production plant. The samples of natural zooplankton were collected from April/May to the end of June during two consecutive spring blooms. For comparison, the same vitamins were measured in nauplii of Artemia franciscana, just after hatching and after enrichment for 24 and 48 h with a commercial diet. The changes in vitamin content in zooplankton, which consisted mainly of Temora longicornis, during the two spring blooms were <2.5-fold for each specific vitamin. The large number of high quality halibut fry produced using the plankton as a dietary source suggests that the plankton contained sufficient amounts of all water-soluble vitamins. With the exception of thiamin and ascorbic acid, Artemia– both newly hatched and enriched for 24 and 48 h – contained higher levels of water-soluble vitamins than natural zooplankton. Enrichment feeding of Artemia for 48 h stabilised or increased the level of the water-soluble vitamins. Received: 8 March 1999 / Accepted: 21 February 2000     

Continuous plankton records: Geographical variations in numerical abundance,biomass and production of euphausiids in the North Atlantic Ocean and the North Sea

Geographical variations in the numbers, biomass and production of euphausiids and the contribution of common species to the total are described from samples taken during 1966 and 1967 in the North Atlantic Ocean and the North Sea by the Continuous Plankton Recorder at 10 m depth. Euphausiids were most abundant in the central and western North Atlantic Ocean and the Norwegian Sea. Thysanoessa longicaudata (Krøyer) was numerically dominant. Biomass was greatest in the Norwegian Sea and the north-eastern North Sea where Meganyctiphanes norvegica (M. Sars) accounted for 81 and 59%, respectively, of the total biomass. Production was highest off Nova Scotia and in Iberian coastal waters; the dominant species were T. raschi (M. Sars) in the former area and Nyctiphanes couchi (Bell) in the latter. The mean P:B ratios were correlated with temperature.     

Growth and herbivory by heterotrophic dinoflagellates in the Southern Ocean,studied by microcosm experiments

Growth and herbivory of heterotrophic dinoflagellates (Gymnodinium sp.) from the Weddell Sea and the Weddell/Scotia Confluence were studied in 1988 in 100-liter microcosms. The microcosms were screened through 200-µm or 20-µm mesh nets and incubated for 12 d at 1 °C under artificial light. Mean cell volume of dinoflagellates was 1 000 to 1 500µm³, and that of their phytoplankton prey 360 to 430µm³. Dinoflagellate growth rate followed a Holling type II functional response, with a maximum growth rate of 0.3 d^–1 and half-saturation food concentrations of 1.0µg chlorophylla l^–1, 50µg C l^–1, or 1 500 cells ml^–1. Carbon budgets based on¹⁴CO₂ assimilation and biomasses of phytoplankton and heterotrophic dinoflagellates suggested a balance between phytoplankton grazing loss and dinoflagellate consumption, assuming a dinoflagellate carbon conversion efficiency of 40%. Applying this to the functional response yielded estimates of maximum ingestion rate (0.8µg Cµg^–1 C d^–1, or 6 pg C dinoflagellate^–1 h^–1) and maximum clearance (0.8 to 1.2 × 10⁵ body volumes h^–1, or 80 to 120 nl ind.^–1 h^–1). The microcosm experiments suggested that heterotrophic dinoflagellates may contribute significantly to maintenance of low phytoplankton biomass in the Southern Ocean.     

Potential impact of climate warming on the recruitment of an economically and ecologically important species, the European lobster (Homarus gammarus) at Helgoland, North Sea

A laboratory-based study was performed to assess the impact of climate warming on the recruitment of the endangered population of the European lobster (Homarus gammarus) at Helgoland (North Sea, German Bight). Egg-bearing females collected in situ just after spawning in late summer were subjected to various seasonal temperature regimes. Regimes with elevated temperatures (mild winters) resulted in a strong seasonal forward shift of larval hatching. Hatching took place at significantly lower temperatures than under regimes with normal winters. Experiments on larval development across a range of constant temperatures showed that no successful larval development occurred at temperatures below 14°C. Larval survival increased from 9% at 14°C to 80% at 22°C, while duration of larval development decreased correspondingly from 26 to 13 days. We hypothesize that an ongoing warming of the North Sea will strongly affect the recruitment success of the Helgoland lobster, mainly resulting from a decoupling of the seasonal peak appearance of larvae from optimal external conditions (temperature, food availability) for larval development.     

The annual cycle of heterotrophic planktonic ciliates in the waters surrounding the Isles of Shoals,Gulf of Maine: an assessment of their trophic role

Macrofauna living on subtidal rocks reefs in southern California excrete ammonium, a potentially important nutrient for benthic algae. Ammonium excretion rates of eleven macroinvertebrate and five fish taxa were determined from a total of 324 in situ incubations conducted between October 1984 and August 1985 at 14 to 17 m depths off Santa Catalina Island, California. Total ammonium excretion ranged from over 100 mol h^-1 by the kelp bass Paralabrax clathratus to less than 0.1 mol h^-1 by the gastropod Conus californicus. Weight-specific ammonium excretion generally ranged from 0.5 to 4 mol g^-1 h^-1 in invertebrates and from 3 to 7 mol g^-1 h^-1 in fishes. Intraspecific excretion rates varied substantially. Coefficient of variation of excretion rates were higher than reported for laboratory studies and multiple regression indicated that 50 to 90% of the variation in ammonium excretion rates of five species studied in detail could not be explained by the combined variation in dry weight, water temperature, time of day, and incubation dates. The excretion data, along with estimates of population densities and size-frequency distributions, indicate that benthic macrofauna release a total of 25 to 30 mol NH ₄ ⁺ m^-2 h^-1 both day and night. The species that generally make the largest contributions are a gobiid fish (Lythrypnus dalli), followed by three gastropods (Astraea undosa, Tegula eiseni, and T. aureotincta) and a sea urchin (Centrostephanus coronatus). The amount of ammonium excreted by these macrofauna on rocky reefs is insignificant compared to our previously published data on the nighttime excretion of blacksmith (Chromis punctipinnis), a pomacentrid fish that feeds in the water column during the day and shelters on the reef at night. Including blacksmiths, we estimate that the amount released by rocky-reef macrofauna at night is >280 mol m^-2 h^-1, a rate that is similar to that for many other marine communities. Additional studies are required to determine if benthic algae utilize ammonium released by these macrofauna, especially at night.Contribution No. 58 of the Ocean Studies Institute; Contribution No. 123 of the Catalina Marine Science Center     

Species-specific grazing rates of heterotrophic dinoflagellates in oceanic waters,measured with a dual-label radioisotope technique

A dual-isotope method was developed to measure grazing rates and food preferences of individual species of heterotrophic dinoflagellates from natural populations, collected from the Slope, Gulf Stream, and Sargasso Sea and from a transect from Iceland to New England, in 1983. The isotope method measures the grazing rates of microzooplankton which cannot be separated in natural populations on the basis of size. Tritiated-thymidine and ¹⁴C-bicarbonate were used to label natural heterotrophic and autotrophic food, respectively. Nine oceanic dinoflagellate species in the genera Protoperidinium, Podolampas, and Diplopsalis fed on both heterotrophic and autotrophic food particles with clearance rates of 0.4 to 8.0 l cell^-1 h^-1, based on ³H incorporation, and 0.0 to 28.3 l cell^-1 h^-1, based on ¹⁴C incorporation. Two dinoflagellate species, Protoperidinium ovatum and Podolampas palmipes, fed only on ³H-labelled food particles. Several species of dinoflagellates fed on bacteria (<1 m) which had been prelabelled with ³H-thymidine. The clearance rates of heterotrophic dinoflagellates and ciliates were similar and within the range of tintinnid ciliate clearance rates reported in the literature. As heterotrophic dinoflagellates and ciliates can have comparable abundances in oceanic waters, we conclude that heterotrophic dinoflagellates may have an equally important impact as microheterotrophic grazers of phytoplankton and bacteria in oceanic waters.Partially supported by a grant from the National Science Foundation, OCE-81-17744     

Variability in dry weight and vertical distributions of decapod larvae in the Irish Sea and North Sea during the spring

Accumulation of waterborne cadmium in Littorina littorea, Mytilus edulis and Carcinus maenas (collected in 1988 and 1989 around the island of Funen, Denmark) was investigated in a matrix of salinities (10 to 30) and calcium concentrations (2.9 to 8.9 mM Ca⁺⁺). Cadmium accumulation rates in soft parts of L. littorina, soft parts and shells of M. edulis and whole bodies and exoskeletons of C. maenas decreased with increasing salinity. Changes in the calcium concentrations accounted for 72% of the salinity effect on cadmium accumulation rates in L. littorina, whereas calcium concentrations had little or no effect on cadmium accumulation in M. edulis. Cadmium accumulation in the whole body of C. maenas was affected equally by calcium concentrations and total salinity, whereas accumulation in the exoskeleton was mainly affected by changes in total salinity. Individual variability in cadmium accumulation in the organs of C. maenas was greater than the variation attributable either to changes in ambient calcium concentrations or total salinity. An appreciable amount of the inter-individual variability in the cadmium accumulation in all three species was correlated with wet:dry weight ratios of the tissues and size of the organisms.     

Biomass partitioning of benthic microbes in a Baltic inlet: relationships between bacteria, algae, heterotrophic flagellates and ciliates

 The structure of a benthic microbial food web and its seasonal changes were studied in the shallow brackish waters of the island of Hiddensee, northeastern Germany, at two sites in close proximity by monthly or bimonthly sampling from July 1995 to June 1996. Abundance and biomass of phototrophic and non-phototrophic bacteria, heterotrophic flagellates (HF) and ciliates as well as the biomass of microphytobenthos were determined in the upper 0.3 cm sediment layer. Abundance of organisms showed strong positive correlation with water temperature, with the exception of the bacteria. Non-phototrophic bacterial numbers ranged from 7 × 10⁸ to 6.7 × 10⁹ cells cm⁻³ and phototrophic bacterial abundance from 4 × 10⁷ to 2.7 × 10⁸. Heterotrophic protist abundance ranged from 8 × 10³ to 104 × 10³ ind cm⁻³ for HF and from 39 to 747 ind cm⁻³ for ciliates. The biomass partitioning demonstrated the primary importance of non-phototrophic bacteria (min. 0.83, max. 84.87 μg C cm⁻³), followed by the microphytobenthos (min. 1.32, max. 50.93 μg C cm⁻³). The heterotrophic protists contributed roughly the same fraction to the total microbial biomass, with the biomass of the HF being slightly higher (HF 0.23 to 1.76 μg C cm⁻³, ciliates 0.04 to 1.17 μg C cm⁻³). Taxonomic classification of the benthic HF revealed the euglenids to be the most important group in terms of abundance and biomass, followed by thaumatomastigids and kinetoplastids. Other important groups were apusomonads, cercomonads, pedinellids and cryptomonads. The structure of the HF assemblage showed strong seasonal changes with euglenids being the most abundant taxa in summer, while apusomonads and thaumatomastigids were predominant in winter. Similar to the pelagic microbial food web, benthic picophototrophic bacteria were occasionally abundant, and the feeding modes of heterotrophic protists exhibited a great variety (predominantly omnivores, bacterivores, herbivores or predators). Filter-feeding HF were of little importance. Contrary to the pelagic environment, a top-down control on total benthic bacterial numbers by HF seemed unlikely at the studied stations which were characterised by muddy sand. Received: 6 January 1999 / Accepted: 21 October 1999     

Microphagous ciliates in mesohaline Chesapeake Bay waters: Estimates of growth rates and consumption by copepods

Growth rates of microphagous ciliates (forms which feed primarily on picoplankton-sized prey) were estimated, along with rates of their consumption by copepods, in shipboard experiments conducted in the mesohaline portion of Chesapeake Bay, USA, under contrasting water column conditions in April, June, and August 1987. Estimates were based on temporal changes in cell densities in size-fractionated water samples incubated under in situ conditions. In April, at low temperatures (7 to 10°C) and with oxygen present throughout the water column, similar generation times of ca. 1 to 1.5 d were estimated for surface and deep water (24 m) ciliate populations. In June, water was anoxic below 12 m and a distinct anoxic microphage community grew at about twice the rate of the surface community with generation times of ca. 7 and 14 h, respectively. In August, bottom water was again anoxic, but the sameStrobilidium sp. dominated both surface and deep waters with low or no growth apparent in anoxic waters and a generation time of ca. 8 h in surface waters. Copepod (primarilyAcartia tonsa Dana nauplii) clearance rates for microphagous ciliates in surface waters were 0.11, 0.56, and 0.53 ml h^–1 copepod^–1 for April, June and August, respectively. Calculation of removal rates, based on average densities, indicated that from 34 to 200% of surface waters were cleared d^–1 of microphagous ciliates by copepods.     

Prey size selection,feeding rates and growth dynamics of heterotrophic dinoflagellates with special emphasis on Gyrodinium spirale

The relationship between size and growth rate in heterotrophic dinoflagellates (collected from the Kattegat, Denmark during 1989 and 1990) was studied. In addition, prey size selection, feeding rates and growth dynamics were studied for the naked heterotrophic dinoflagellate Gyrodinium spirale Bergh. Heterotrophic dinoflagellates have growth rates which are approximately three times lower than that of their potential competitors, the ciliates. G. spirale requires a relatively high prey concentration in order to grow. Ingestion rate at the maintenance level is about half of that of maximum ingestion rate. Consequently, yield is lower than typically found for planktonic protozoa. When exposed to low prey concentrations, the dinoflagellate is able to reduce its metabolism and thus prolong survival. The optimum prey particle size for G. spirale, which feeds by direct engulfment, corresponds to its own size. The ability to ingest relatively large prey may explain why these organisms are competitive in nature.