The effects of fluctuations in North Sea flows on zooplankton abundance

Time series of annual mean zooplankton abundance observed bythe Continuous Plankton Recorder Survey in the North Sea duringthe last three decades are compared with transports from a depth-averagedstorm-surge model by means of multiple regression analysis.In the northern areas, more than half the variance of Calanusfinmarchicus can be accounted for by the winter flows. Thereare no statistically significant relationships in other NorthSea areas or by using the flows from other seasons. This isconsistent with the accepted view that this copepod overwintersin the North Atlantic and enters the North Sea in the earlyspring. The model flows are strongly related to the strengthof the North Atlantic Oscillation (NAO). which may explain therecently reported association of C.finmarchicus with the NAO.Other species do not exhibit statistically significant relationshipswith any of the model flows.     

Tracking seasonal changes in North Sea zooplankton trophic dynamics using stable isotopes

Trophodynamics of meso-zooplankton in the North Sea (NS) were assessed at a site in the southern NS, and at a shallow and a deep site in the central NS. Offshore and neritic species from different ecological niches, including Calanus spp., Temora spp. and Sagitta spp., were collected during seven cruises over 14 months from 2007 to 2008. Bulk stable isotope (SI) analysis, phospholipid-derived fatty acid (PLFA) compositions, and δ ¹³C_PLFA data of meso-zooplankton and particulate organic matter (POM) were used to describe changes in zooplankton relative trophic positions (RTPs) and trophodynamics. The aim of the study was to test the hypothesis that the RTPs of zooplankton in the North Sea vary spatially and seasonally, in response to hydrographic variability, with the microbial food web playing an important role at times. Zooplankton RTPs tended to be higher during winter and lower during the phytoplankton bloom in spring. RTPs were highest for predators such as Sagitta sp. and Calanus helgolandicus and lowest for small copepods such as Pseudocalanus elongatus and zoea larvae (Brachyura). δ ¹⁵N_POM-based RTPs were only moderate surrogates for animals’ ecological niches, because of the plasticity in source materials from the herbivorous and the microbial loop food web. Common (16:0) and essential (eicosapentaenoic acid, EPA and docosahexaenoic acid, DHA) structural lipids showed relatively constant abundances. This could be explained by incorporation of PLFAs with δ ¹³C signatures which followed seasonal changes in bulk δ ¹³C_POM and PLFA δ ¹³C_POM signatures. This study highlighted the complementarity of three biogeochemical approaches for trophodynamic studies and substantiated conceptual views of size-based food web analysis, in which small individuals of large species may be functionally equivalent to large individuals of small species. Seasonal and spatial variability was also important in altering the relative importance of the herbivorous and microbial food webs.     

An investigation into the zooplankton composition of a prominent 38-kHz scattering layer in the North Sea

This study aims to determine the contribution made by zooplanktonto acoustic scattering layers, which are particularly strongat 38 kHz, in the northern North Sea in summer. It uses a combinationof net sampling and forward and inverse acoustic modelling ofdata collected at 38, 120 and 200 kHz in July 2003. Zooplanktonsamples were collected from regions of strong acoustic scatteringin depths to 50 m, using a undulating towed (U-tow) vehicle.Acoustic data recorded simultaneously were scrutinized to determineactual backscattering, expressed as mean volume backscatteringstrength (MVBS) (dB). This observed MVBS (MVBS_obs) was comparedwith backscattering predicted by application of appropriateacoustic models (MVBS_pred) to sampled densities of zooplankton.In all instances, MVBS_obs was greater than MVBS_pred, with thedifference considerably more pronounced at 38 kHz. There wasa weak correlation between MVBS_pred and MVBS_obs at all threefrequencies, with the greatest correlation (r = 0.450, P = 0.545)at 120 kHz. A number of candidate acoustic models were invertedin order to infer the most likely type of scatterer. In mostcases, scatterers with a gas inclusion were predicted by thismethod. Potential sources of inconsistencies between MVBS_predand MVBS_obs were identified which, when considered alongsidethe presented forward and inverse solutions, indicate that 38kHz scattering in particular must be due to sources other thansampled zooplankton.     

Aspects of Scotia Sea zooplankton

Information on small scale distributions of three species of Antarctic zooplankton is reviewed. Aggregations of the euphausiid Euphausia superba , the tunicate Salpa thompsoni , and the amphipod Parathemisto gaudichaudii are compared, and the manner in which such aggregations mav arise is discussed. A possible relationship between swarming and feeding activity in E. superba is suggested in which krill are thought to be dispersed whilst feeding but that on repletion they swarm. It is thought that this may account for this species' irregular spatial distribution as recorded bv previous expeditions. A further consequence of this theory is that during the Winter swarming will be minimal.     

渤海浮游动物群落生态特点Ⅲ.部分浮游动物数量分布和季节变动

用1959年全国海洋普查中网浮游动物标本,分析渤海水母类,枝角类,糠虾,毛虾和幼虫类的数量分布和季节变化,结果表明,水母类以近岸低盐种为主,主要出现在莱州湾,渤海湾和辽东湾近岸水域,冬季出现的种类较少,优势种为八斑芮氏水母;夏季种类较多,优势种为半球杯水母和嵊山多胃水母等,鸟喙尖头蚤为近岸高温低盐种,主要出现在莱州湾,数量在6月份达到最大值,箭虫是海主要肉食性种类之一,其数量高峰期出现在9月份,长住囊虫的季节变化呈双峰型变化,第1次数量高峰出现在5月份,第2次数量高峰出现在8月份。该种大部分时间主要分布在莱州湾,渤海湾和辽东湾近岸水域,渤海出现的糠虾有长额刺糠虾和漂浮囊糠虾,其中长额否则糠虾受温度的影响较小,在春,夏秋,各有一个数量高峰;漂浮囊糠虾在秋末数量明显增加,这两种糠虾主要分布在渤海湾和辽东湾近岸水域。／三叶针尾涟虫属于低温种类,主要出现在秋季和冬季,细长脚绒属夏季高温种,在6,7月份数量较高,该种的分布受盐度影响比较大,主要出现在受黄海海流影响的谪盐水域,数量较大的幼虫有桡足类的六肢幼体,多毛类海稚虫科的幼体,以壳类幼体和腹足类幼体,其中桡足类六肢幼体和海稚虫科的幼体在5月份出现较多,腹足类幼体在6．7月份数量较高,双壳类幼体在8月份数量较高。     

Long-term dynamics of zooplankton in the southeastern Baltic Sea

Long-term research in the Baltic Sea revealed the basic trends of zooplankton community variations depending on oceanographic processes. Alternation of the periods of increase and decrease in salinity of the Baltic Sea against the background of climate changes (temperature increase) and eutrophication affect the state of the entire Baltic ecosystem, including zooplankton. For these periods, the dynamics of zooplankton in the Baltic Sea were analyzed based on literature data and results of regular research in the southeastern Baltic Sea during 1998–2007. The changes in the hydrological situation were accompanied by significant changes in the zooplankton community. In the 1990s–2000s, the abundance and biomass of brackish-water and thermophilous species primarily of Cladocera and Copepoda increased markedly. The role of the previously dominant marine copepod Pseudocalanus elongatus decreased due to salinity reduction in the deep-water part of the Baltic Sea. Maximum development of zooplankton occurred in years of the greatest warming-up of the water (2001, 2005–2007) against the background of a general positive trend of zooplankton abundance in the last decade.     

渤海浮游动物群落生态特点 Ⅱ.桡足类数量分布及变动

用1959年全国海洋普查浮游动物中网样品分析渤海浮游动物。尤其是中小型浮游动物的数量分布及变动特点。结果表明,渤海浮游动物全年平均丰度为3841ind／m^3,最大33756ind/m^3,最小6．5ind／m^3。其中桡足类年平均密度为3413ind／m^3,是数量的主要构成者。渤海浮游动物主要的高峰期出现在夏季,此外春、秋两季各自有一个小的高峰。春季峰主要是由双毛纺锤水蚤(Acartia bifilosa)的大量繁殖造成,秋季峰主要是由拟长腹剑水蚤(Oithona similis)构成。夏季峰是由于小拟哲水蚤(Paraclanus parvus)、强额拟哲水蚤(Paracalanus crassirostris)、拟长腹剑水蚤等多种桡足类和一些幼虫大量繁殖造成。渤海浮游动物季节演替比较明显。双毛纺锤水蚤、墨氏胸刺水蚤(Centropages mcmmurrichi)和中华哲水蚤(Calanus sinicus)是春季的优势种;小拟哲水蚤、拟长腹剑水蚤、近缘大眼剑水蚤(Corycaeus affinis)和太平洋纺锤水蚤（Acartia pacifica)是夏季的优势种;拟长腹剑水蚤、真刺唇角水蚤（Lubidocera euchaeta)等是秋季的优势种。数量的水平分布同温度的水平分布趋势基本一致。同盐度趋势相反。     

Interannual changes in the biomass of the Black Sea gelatinous zooplankton

The 15 year changes in the total gelatinous biomass consistof a general trend to increase, from 250 gm^–2 in 1980to 2500 g m^–2 in 1995, and fluctuations with a periodof –4 years performed against the background of this trendin the upper 150 m layer. Different species occupying distincttrophic niches form these peaks. When represented as the percentageof the total zooplankton biomass, Aurelia aurita and Mnemiopsisleidyi exhibit the contra-phase fluctuations where a sharp increasein M.leidyi is accompanied by a decrease in the A.aurita biornass.     

Mangrove zooplankton of North Queensland,Australia

Measurements of plankton community structure and trophic resourcespotentially available to planktonic copepods were made in the mangroveestuaries of six rivers in Northeastern Australia. The Pascoe, Claudie,Lockhart, McIvor and Daintree Rivers represent wet tropical systems on CapeYork, whereas the Haughton River estuary has restricted freshwater inflowbecause of a drier climate and freshwater diversion for agriculture. TheHaughton River was sampled approximately monthly between October 1992 andMay 1994, and had a mean abundance of zooplankton >37 μm of 200l⁻¹ (range 60–500 l⁻¹). The Cape Yorkrivers were sampled infrequently, and zooplankton abundances ranged between0.4 and 1400 l⁻¹. The zooplankton of all rivers was dominatedby copepods, particularly representatives of the genus Oithona which werecharacteristic of a distinct mangrove fauna. Physical forcing influencedthe zooplankton of mangrove estuaries much more than the measured biologicalvariables. The water column was characterised by high concentrations ofparticulate matter, up to 3.3 mg l⁻¹ C and 1.1 mgl⁻¹ N, of low food quality (as indicated by the C:N ratio).Phytoplankton biomass (as chlorophyll a) in all six rivers was on averagefour-fold greater than in neighbouring coastal waters (1.1–12.6μg l⁻¹), and 25% of this chlorophyll a wasderived from cells >10 μm, and thus potentially available tocopepods. The degree of mixing, determined by the combination of tidal stateand the extent of freshwater input, appears to drive both the quantity andquality of particulate material available to higher consumers and thedistribution of zooplankton communities within mangrove estuaries. This revised version was published online in July 2006 with corrections to the Cover Date.     

Mangrove zooplankton of North Queensland,Australia

Food consumption, growth, fish length distributions,population sizes and habitat use of the salmonids intwo lakes in the Høylandet area were studied in1986–89. The allopatric brown trout (Salmotrutta L.) in the tarn Røyrtjønna (27 ha) fed mainlyon organisms at the lake surface , crustaceanplankton, Trichoptera and Chironomidae. Only 5% ofthe trout reached an age of 6 years and a length of25 cm. Sexual maturation started at age 3 and a lengthof 14 cm. Through mark – recapture technique thenumber of trout >10 cm was estimated to 115 ha^-1.Growth, fish length frequencies and sexualmaturation of the sympatric brown trout and Arcticcharr (Salvelinus alpinus (L.)) in LakeStorgrønningen (530 ha) were not much different. TheStorgrønningen charr fed chiefly on zooplankton whichby volume represented 33% for the trout. The foodconsumption of Storgrønningen trout was at maximum inJuly with 2.06 mg food (d.w.) per g live fish and forcharr in September with 1.26 mg food. The maximumsize-independent growth rate of trout was 5.2%day^-1 in late June, and for charr 4.1%day^-1 in late July. Seventy percent of theirseasonal growth took place before 15 August. The charrstayed mainly deeper than 3-4 m, at water temperatures<15 °C. Brown trout stayed mainly the littoralzone and in near surface water of the pelagic. Thenumber of pelagic charr was estimated hydroacusticallyto 50 ind. ha^-1. The charr spawn in thelake. Mean numbers of juvenile trout in the twolargest tributaries were 26 and 48 per 100 m².Their annual length increment was 2.8–3.4 cm. Noindication of acidification or other human inducedimpacts were found. The lakes and their tributariesrepresent complex aquatic systems, representative forpristine oligotrophic Norwegian lowland lakes.John W. Jensen died shortly after easter in 1996     

Checklist of fauna found in zooplankton samples from the Barents Sea

We present a list of zooplankton taxa occurring in the Barents Sea based on our analysis of 360 samples from the upper 100 m of the water column as well as on reports from published sources. Moreover, we provide information on adult size, ecological and geographical distribution, habitat preference and frequency of occurrence of each taxon. The list encompasses a total of 374 species and higher taxa. Most are mesozooplankton, with Copepoda prevailing (153 species), and belong to the holoplankton. However, pseudoplankton, organisms only circumstantially occurring in the water column, also comprise a significant part of the fauna (86 taxa). The coastal zooplankton (264 neritic species) was richer than the fauna of the open sea. In our samples, the majority of the zooplankton taxa were rarely found (94 of 157 taxa, equivalent of 60%), 46 taxa (29%) were moderate and only 17 taxa (11%) were common (found in >80% of the samples). Among these, the copepods Calanus finmarchicus, C. glacialis, Oithona similis and Pseudocalanus minutus were most frequent.     

Temporal and spatial variation of coastal zooplankton in the Baltic Sea

When analysing temporal variation monthly mean abundances of zooplankton sampled at a eoastal station in the northern Baltic Sea between 1976 and 1988 showed the lowest between-year variation in the summers The coefficients of variation were estimated at 50% for rotifers in June. 70% for cladocerans in August and between 30 and 50% for different copepodite stages in August Moreover in the summers, all dominating zooplankton groups were abundant Estimates of yearly biomass or production of zooplankton must, however, be based on frequent sampling during the whole year because of large differences between months in zooplankton composition and abundance
In 1989. spatial variation was analyzed by sampling zooplankton on three occasions along a 9 8 km transect with 15 stations, 700 m apart All taxonomic/life-stage groupings of zooplankton were heterogeneously distributed Geostatistical analysis did not give a definite answer to what the minimum distance between stations should be to obtain independent samples but indicated that 700 to 1400 m would be sufficient in most cases For most of the zooplankton groupings the coefficients of variation in the sampled area were estimated at 20-40% on all sampling occasions     

Zooplankton research in the North Sea

Summary Water types of the North Sea with different plankton are the thermally stratified northern and central regions with a relatively nutrient rich inflow of Atlantic water, a mixed region in the southern North Sea with a poor inflow of Channel water, and turbid narrow coastal zones with inflow of nutrient rich river water. Plankton studies reveal that the primary production starts early, in February, in the southern region, but is delayed in the coastal zones by turbidity. In stratified areas the algal spring bloom is delayed by mixing towards the greater depth and usually starts with the onset of thermal stratification. The spring bloom soon declines and the algae remain on a low density level in summer, presumably due to depletion of nutrients in the euphotic zone. The coastal zones and the frontal zones between mixed and stratified water have a relatively high summer primary production.The herbivores (mainly planktonic copepodes and the tunicateOikopleura dioica) grow and increase in number when the temperature rises and food is available. There is a considerable mismatch with the algal spring bloom, which comes too early and is too short in most regions. The best coincidence occurs in the coastal zones and maybe the frontal zones. Carnivores build high biomasses in late summer and fall in the coastal regions and compete with fish larvae for food and also kill many fish larvae. The large scyphomedusae are most important in this respect.The overall yearly primary production of the North Sea is estimated to be about 100 mg C.m^–2. The estimates for herbivores and fish are 20 and 1 mg C.m^–2. Considering a growth efficiency of 20%, the herbivores must consume all algae produced. The indication of a low consumption due to bad phenological coincidence in most regions leads to the assumption, that either primary production is underestimated or there is a considerable influx of organic matter from the Atlantic Ocean. During June–July 1979 the carnivore consumption was estimated in the coastal zone of the Southern Bight to be 39 mg C.m^–2.d^–1 at a copepod production of 20 mg C.m^–2.d^–1. Consumption by fish larvae and large jellyfish (Cyanea lamarckii) was 15% and 74%, respectively.It seems clear that the productivity of the North Sea depends highly on coastal and frontal zones, where herbivores find sufficient food at optimal growth conditions. Most organic matter will at the end be consumed by invertebrate carnivores, which urge fish populations to reproduce early in spring or to recruit at remote places.     

Bacteriophage Diversity in the North Sea

In recent years interest in bacteriophages in aquatic environments has increased. Electron microscopy studies have revealed high numbers of phage particles (10⁴ to 10⁷ particles per ml) in the marine environment. However, the ecological role of these bacteriophages is still unknown, and the role of the phages in the control of bacterioplankton by lysis and the potential for gene transfer are disputed. Even the basic questions of the genetic relationships of the phages and the diversity of phage-host systems in aquatic environments have not been answered. We investigated the diversity of 22 phage-host systems after 85 phages were collected at one station near a German island, Helgoland, located in the North Sea. The relationships among the phages were determined by electron microscopy, DNA-DNA hybridization, and host range studies. On the basis of morphology, 11 phages were assigned to the virus family Myoviridae, 7 phages were assigned to the family Siphoviridae, and 4 phages were assigned to the family Podoviridae. DNA-DNA hybridization confirmed that there was no DNA homology between phages belonging to different families. We found that the 22 marine bacteriophages belonged to 13 different species. The host bacteria were differentiated by morphological and physiological tests and by 16S ribosomal DNA sequencing. All of the bacteria were gram negative, facultatively anaerobic, motile, and coccoid. The 16S rRNA sequences of the bacteria exhibited high levels of similarity (98 to 99%) with the sequences of organisms belonging to the genus Pseudoalteromonas, which belongs to the γ subdivision of the class Proteobacteria.The marine bacterial community is responsible for a considerable portion of primary production and regeneration of nutrients in the microbial loop and is associated with a great variety of marine bacteriophages (5, 12). These phages are capable of infecting a large portion of the bacterioplankton (32, 34). It is assumed that as part of the marine food web, bacteriophages play important quantitative and qualitative roles in controlling marine bacterial populations (8, 24, 34, 39, 45). The phenotypic diversity and genotypic diversity of the phage populations are related to the interaction between phages and their host organisms, which provides a tool for understanding the interaction itself (13). To estimate the influence of marine bacteriophages on the diversity of bacterioplankton, we investigated phage diversity. The virus species concept proposed by Murphy et al. (37) delineates seven different families of bacteriophages based on morphological criteria and provides criteria for new phage species based on several traits, such as DNA homologies, serological data, protein profiles, and host ranges.In this paper, we describe the diversity and genetic relationships of marine phages based on investigations of 22 representatives from 85 phage-host systems (35, 36) collected between 1988 and 1992 from waters around an island, Helgoland, located in the North Sea. All of the phages were virulent and formed plaques on their host bacteria. We assigned the phages to different virus families, species, and strains based on morphology, DNA homology, and host range. Furthermore, we characterized the phenotypic and genotypic features of the host bacteria.     

A decline in primary production in the North Sea over 25 years,associated with reductions in zooplankton abundance and fish stock recruitment

Phytoplankton primary production is at the base of the marine food web; changes in primary production have direct or indirect effects on higher trophic levels, from zooplankton organisms to marine mammals and seabirds. Here, we present a new time‐series on gross primary production in the North Sea, from 1988 to 2013, estimated using in situ measurements of chlorophyll and underwater light. This shows that recent decades have seen a significant decline in primary production in the North Sea. Moreover, primary production differs in magnitude between six hydrodynamic regions within the North Sea. Sea surface warming and reduced riverine nutrient inputs are found to be likely contributors to the declining levels of primary production. In turn, significant correlations are found between observed changes in primary production and the dynamics of higher trophic levels including (small) copepods and a standardized index of fish recruitment, averaged over seven stocks of high commercial significance in the North Sea. Given positive (bottom‐up) associations between primary production, zooplankton abundance and fish stock recruitment, this study provides strong evidence that if the decline in primary production continues, knock‐on effects upon the productivity of fisheries are to be expected unless these fisheries are managed effectively and cautiously.     

Distribution of capsulated bacterioplankton in the North Atlantic and North Sea

In laboratory experiments, bacterioplankton were incubated under different nutrient conditions, and the percentage of bacteria exhibiting a polysaccharidic capsule (capsulated bacteria) and that of CTC (cyanotetrazolium chloride)-positive and therefore metabolically highly active bacteria were determined. In these seawater cultures amended with nutrients more than 95% of the CTC-positive cells exhibited a capsule. During two cruises, one to the North Atlantic and one to the North Sea, we investigated the distribution of capsulated bacteria throughout the water column. Capsulated bacteria were generally more abundant in eutrophic surface waters than in deeper layers or more oligotrophic regions. In the upper 100 m of the North Atlantic, about 6–14% of the total bacterioplankton community was capsulated, while in the layers below 100 m depth, 97% of the bacteria lacked a visible capsule. The percentage of capsulated bacteria correlated with bacterial abundance and production, and chlorophyll a concentration. Also, the bioavailability of DOC (dissolved organic carbon), estimated by the ratio between bacterial production and DOC concentration, significantly correlated with the percentage of capsulated bacteria. In the North Sea, the contribution of capsulated bacteria to the total number of bacteria decreased from the surface (3 m depth) to the near-bottom (25–35 m) layers from 20% to 14% capsulated bacteria. In the nearshore area of the North Sea, about 27% of the bacteria exhibited a capsule. Overall, a pronounced decrease in the contribution of capsulated bacteria to the total bacterial abundance was detectable from the eutrophic coastal environment to the open North Atlantic. Using this epifluorescence-based technique to enumerate capsulated bacterioplankton thus allowed us to routinely assess the number of capsulated bacteria even in the oceanic water column. Based on the data obtained in this study we conclude that almost all metabolically highly active bacteria exhibit a capsule, but also some of the metabolically less active cells express a polysaccharide capsule detectable with this method.     

Euthanasia across the North Sea

The United Kingdom and The Netherlands are separated by a narrow stretch of sea but in terms of an understanding of euthanasia they seem to be light years apart. An attempt to bridge the information gap seems in order.The position paper on euthanasia published by the Royal Dutch Medical Association contains 16 typed pages.¹ In contrast, the report on euthanasia issued by a working party of the British Medical Association is 80 pages in small print.²     

Benthic community structures in the North Sea

Coherent assemblages of marine benthic species have been recognized from the early twentieth century, and the classical papers of Petersen (1914, 1918) were based on studies of limited areas in the North Sea. In 1986, a synoptic survey of the North Sea north to 57°N was undertaken by a group of ten laboratories from seven North Sea countries. The results of this survey have recently been published (Heip et al., 1992a, b; Künitzer et al., 1992; Huys et al., 1992), and some of the results are summarized in this paper. The analysis of the macrofauna is based on slightly more than 700 taxa. In general, the North Sea macrofauna consists of northern species extending south to the northern margins of the Dogger Bank, and southern species extending north to the 100 m depth line. The central North Sea is an area of overlap of southern and northern species, especially around the 70 m depth contour. Consistent groupings of species are recognized that were summarized in seven faunal groupings. Macrofaunal body weight, density and diversity increase linearly towards the north. Macrofaunal biomass for the whole area averages 7 g adwt. m⁻² and decreases from south to north. Distribution patterns and trends within the meiofauna were very different. Nematodes, which are the dominant taxon overall, are least abundant in the sandy sediments of the Southern Bight, then increase to a maximum around 53° 30′ N and slowly decrease again towards the north. Copepod density and diversity are highest in the Southern Bight, due to the presence of many interstitial species. A large number of species new to science were recorded by the North Sea Benthos Survey and about 1500 species are expected to occur. Copepods show very distinct assemblages according to water depth and sediment type. The contrasting patterns in latitudinal gradients of body weight and number of species of macro- and meiofauna can be only partially explained. Latitude and sediment characteristics, such as grain size and content in plant pigments, and water depth, determine part of the variance in species composition, density and biomass of the benthic fauna, but the patterns that are observed are different for different benthic groups, requiring careful consideration as to their use in biological monitoring procedures. Distributions are related to current patterns in the North Sea, annual temperature variations and availability of food. However, large parts of the variance in many parameters remain unexplained.     

Algal production in the west-central North Sea

In 1976, 13 cruises were used for a plankton survey of an areaapproximately 300 x 150 km off the north-east coast of Englandfrom March to November. Results are presented of integratedzooplankton samples, surface measured nitrate, phosphate, silicateand chlorophyll, as well as other environmental characteristics.They show that the spring bloom started from near the DoggerBank and spread to most of the area by the end of April; thisdeclined and a nearshore production zone was seen in summer.This eventually disappeared and an offshore autumnal peak wasfound. The data are interpreted with use of a vertically integratedtwo-dimensional model. The model adequately explains the amplitudeand timing of the two-dimensional distributions of algal standingstock for the first half of the year but provides a poor representationafter that. It is inferred that annual primary production inthis region is low, about 40 g C.m^–2. This is due to lowoverwinter levels of nutrients and a shallow mixed layer depthof water. Pelagic fish are considered to have an ecologicalefficiency of 7–10% and demersal fish of about 4%. Thelatter figure may be due to underexploitation of demersal biomass.