Modelling the basin-scale demography of Calanus finmarchicus in the north-east Atlantic

In this paper, we report on a coupled physical–biological model describing the spatio-temporal distribution of Calanus finmarchicus over an area of the North Atlantic and Norwegian Sea from 56°N, 30°W to 72°N, 20°E. The model, which explicitly represents all the life-history stages, is implemented in a highly efficient discrete space–time format which permits wide-ranging dynamic exploration and parameter optimization. The underlying hydrodynamic driving functions come from the Hamburg Shelf-Ocean Model (HAMSOM). The spatio-temporal distribution of resources powering development and reproduction is inferred from SeaWiFS sea-surface colour observations. We confront the model with distributional data inferred from continuous plankton recorder observations, overwintering distribution data from a variety of EU, UK national and Canadian programmes which were collated as part of the Trans-Atlantic Study of Calanus (TASC) programme, and high-frequency stage-resolved point time-series obtained as part of the TASC programme. We test two competing hypotheses concerning the control of awakening from diapause and conclude that only a mechanism with characteristics similar to photoperiodic control can explain the test data.     

The multinomial logit model: a new tool for exploring Continuous Plankton Recorder data

The use of Generalized Additive Models in the analysis of Continuous Plankton Recorder (CPR) distribution and abundance data for C alanus finmarchicus and C. helgolandicus is demonstrated. Strong emphasis is placed on the influence of long-term trend, seasonality and location. The preponderance of zeros in the data, as well as the particular problems associated with the CPR recording protocol, are adequately handled by the models. Three models are described. In the first two, data within arbitrary spatial compartments are modelled as smooth functions of long-term trend and seasonality, assuming a spatial homogeneity within the area. In the third model, the spatio-temporal distributions of abundances over the North Sea in specific months each year are modelled as a three-dimensional smooth function of long-term trend and location (latitude and longitude). The summaries produced by the models reveal dynamic seasonal, spatial and long-term patterns of abundance, many of which have not been revealed by the previous analytical techniques employed.     

Oil extraction From the Copepod Calanus finmarchicus Using Proteolytic Enzymes

The limited amount of fish oils available has led to extensive search for alternative sources of oils rich in long-chain n-3 polyunsaturated fatty acids (PUFA). The zooplankton Calanus finmarchicus is present in large amounts in the North Atlantic and have lipid-rich stages which can be harvested. The main objective of this study was to investigate if the use of food grade proteolytic enzymes, Alcalase® and Flavourzyme®, could improve oil recovery in an industrial-like process, and to characterize the oil obtained. The results showed a substantially higher oil yield with the use of proteolytic enzymes compared to standard fish oil production technology. The oil from C. finmarchicus had a high content of n-3 PUFA mainly comprised of stearidonic acid (18:4 n-3), eicosapentaenoic acid (EPA), and docosahexaenoic acid (DHA). Wax esters were the dominating lipid class in the oil.     

渤海中华哲水蚤桡足幼体摄食的初步研究

对３４２个含有食物的消化道分析结果表明,中华哲水蚤桡足幼体营浮游植物食性,主要摄食硅藻类,其中辐圆筛藻,未查明的圆筛藻和星脐圆筛藻是它的主要摄食对象对象。它在渤海所食物类群,几科没有季节变化,四季都以硅藻类居绝对优势,食物类群更替率各季平均只有０．５％。它所摄食的食物种类则有一定的季节变化,食种类更替率各季平均为１２．７％。它的摄食强度以冬季为最高,春,夏次之,秋季最低,它对环境中的饵料浮游植物有     

Population dynamics of Calanus chilensis in the Chilean Eastern Boundary Humboldt Current

A study of the population dynamics of Calanus chilensis was carried out in the coastal area of the Mejillones Peninsula (23°S), which is an upwelling site in the Eastern Boundary Humboldt Current. Information was derived from time-series sampling (15 day interval) at three stations, and from two broad-scale cruises: under active upwelling, and under 1997/1998 El Niño conditions. Calanus chilensis showed continuous reproduction throughout the year. In the study area, upwelling takes place intermittently year-round, suggesting that Calanus production is not limited by food. Advection played an important role in structuring the spatial distribution of Calanus , both during upwelling and under El Niño conditions. Population losses from upwelling and offshore transport may be compensated by rapid turnover rates of cohorts, growing at temperature-dependent rates and hence allowing population recovery, even in abnormally warm waters during El Niño conditions.     

Seasonal changes in respiration rates of copepodite stage V Calanus finmarchicus (Gunnerus)

Copepodite stage V Calanus finmarchicus were collected at locations on the continental shelf north of Scotland, in the Faroe–Shetland Channel and west of Ireland on six occasions covering winter, spring and summer from October 1993 to June 1995. Oxygen consumption by the overwintering and active spring/summer population of animals was determined at temperatures close to in situ temperatures. Laboratory measurements of oxygen consumption were also made at standardized temperatures (0°C, 5°C, 7°C and 12.5°C) to determine the sensitivity of animals to temperature change in the different seasons. Rates of oxygen consumption were very low (7–30 μmol O₂ gC⁻¹ h⁻¹) at in situ temperatures during the winter and early spring and significantly higher (105–219 μmol O₂ gC⁻¹ h⁻¹) for the active surface population in May and June. Animals collected from the overwintering population showed no significant response to changes in temperature. Due to the low respiration rates, the calculated rate of decrease in carbon content in diapausing copepodite stage CV was very low (approximately 0.250 μgC day⁻¹). The respiration rates were used to construct a model to estimate survival of the animals with an initial carbon content equivalent to that expected of animals in October. The results showed that in order to survive during winter and have enough energy for moulting and migration to the surface in the spring, these animals have to live at temperatures close to 0°C and be in a diapause state.     

The spring/early summer distribution of Calanus spp. in the northern North Sea and adjacent areas

The spring/early summer distribution of Calanus spp. in the north-eastern Atlantic and the northern North Sea was studied from data collected on three cruises between 1992 and 1995, and compared against estimates from Continuous Plankton Recorder (CPR) data obtained between 1958 and 1993. The patterns observed in both data sets were examined in relation to the hypothesis proposed by Backhaus et al . (1994 ) to explain the annual spring invasion of the northern North Sea by Calanus finmarchicus . The results confirmed that arrival of Calanus spp. in the northern North Sea started in March. The main routes of transport into the area were the Norwegian Trench Atlantic Inflow and East Shetland Atlantic Inflow. The highest abundances of Calanus spp. juveniles (stages CI–IV) were found in May in both CPR and cruise data. Although significant differences in absolute abundance of Calanus spp. were found between CPR and cruise data, similar geographical and temporal patterns of spring advection into the North Sea were derived from both data sources, these patterns being consistent with the hypothesized transport mechanisms.     

两种藻类饲喂中华哲水蚤的繁殖差异

通过室内实验研究了中华哲水蚤摄食三角褐指藻(Phaeodactylumtricornutum)和球等鞭金藻(Isochrysis galbana)后产卵率、孵化率变化。22d的实验时间,摄食三角褐指藻的中华哲水蚤平均产卵率为0.24±(0.07)/d,雌体存活率较低,摄食球等鞭金藻的产卵率为7.84±(1.12)/d,雌体存活亦较高。不论在藻类溶出液和过滤海水中摄食三角褐指藻后所产的卵孵化率都低于摄食球等鞭金藻者。这表明对照球等鞭金藻相比,硅藻三角褐指藻对中华哲水蚤产卵和孵化都存在明显的抑制作用。脂肪酸分析结果显示,在三角褐指藻细胞内,对桡足类繁殖十分重要的多不饱和脂肪酸(PU-FAs)和特定脂肪酸(ω3,ω6)的含量皆较低。     

A Lagrangian ensemble model of Calanus finmarchicus coupled with a 1D ecosystem model

A population dynamics model of Calanus finmarchicus based on Lagrangian particles has been coupled with a 1-D ecosystem model. Each of the particles represents a variable number of copepods which experience the same fate. Therefore all copepods of a single particle represent a cohort and are characterized by a common set of individual properties such as age, development-stage, depth, structural weight (length), lipid pool or food satiation. The physical environment is parameterized by a 1-D-water column with a vertical resolution of 1 m and a maximum depth of 800 m. Copepod food supply is provided by an interactive Eulerian NPZD model where Z represents microzooplankton. The model correctly reproduces both the dynamics of the ecosystem and the life history of the copepods in the Norwegian Sea. Simulated results of trajectories of particles in the water column, and of individual growth and stage development were analysed. Results on seasonal abundance, development time, number of generations, depth profiles, and patterns of diurnal and ontogenic migration are compared with field data from OWS India.     

Measuring copepod abundance in deep‐water winter habitats in the NE Norwegian Sea: intercomparison of results from laser optical plankton counter and multinet

The main purpose of this study was to assess the usefulness of the laser optical plankton counter (LOPC) as a Calanus finmarchicus monitoring tool in the NE Norwegian Sea in winter. To test this, a multinet and an LOPC were used simultaneously to sample overwintering copepods in the Lofoten basin in January 2007. Additional data from an LOPC laboratory experiment were also analyzed to help the interpretation of the field data. Both the laboratory data and the field data indicated that the presence of particles other than zooplankton generally contaminated the zooplankton signal in the LOPC data. However, reliable abundance estimates could be made from the LOPC data by choosing an appropriate size range in the LOPC size distribution. This size range was determined by comparing LOPC abundance estimates from systematically varying size ranges to multinet zooplankton abundance data. The average difference was smallest for a size range of about 900–1500 μm. As C. finmarchicus, stages IV and V, was by far the most abundant species in the NE Norwegian Sea, abundance estimates from the LOPC in this size range were in practice estimates of C. finmarchicus abundance, making the LOPC suitable as a tool for providing high resolution winter data in the NE Norwegian Sea.     

Annual weather variability and its influence on the population dynamics of Calanus finmarchicus

The reaction of the population of Calanus finmarchicus to relatively extreme annual cycles of weather in the North Sea was investigated by one-dimensional model simulations. A population dynamics model for C. finmarchicus was coupled with a physical and a biological upper-layer model for phosphate, phytoplankton and detritus to simulate the development of the successive stages of C. finmarchicus. Observed annual weather cycles were used to drive the physical water column model, the results of which were then input to the plankton model. The simulations yielded the temporal development of all stages of C. finmarchicus over an annual cycle in terms of numbers and weights. Compared with the results reported for 1984 by Carlotti and Radach (1996; Limnol . Oceanogr . 41: 522–539), it appears that the temporal range for the occurrence of the peak concentration of C. finmarchicus may be 2 weeks in the northern North Sea. The simulated variability is in accordance with observations. Years with more continuous primary production such as the year 1984, which was cloudy and windy during the summer, may then provide the optimum conditions with respect to producing an abundance of zooplankton during the following year.     

Covariation in climate, zooplankton biomass and mackerel recruitment in the southern Gulf of St Lawrence

Between 1982 and 1991, an annual survey of stage I egg production of Atlantic mackerel ( Scomber scombrus ) was conducted in June/early July in the southern Gulf of St Lawrence. We investigated the relationship between interannual variability in biomass of zooplankton, determined from the archived survey plankton samples, and mackerel recruitment, estimated from the proportion of three-year-olds in the catch of the commercial fishery. Zooplankton biomass varied by a factor of 2.5, primarily owing to fluctuations in the >1000 μm size fraction. The index of mackerel recruitment fluctuated by a factor of ≈20 and was positively related (linear regression: P  < 0.05; n  = 10) to variations in the zooplankton biomass. Both mackerel recruitment and zooplankton biomass were negatively related (linear regression: P  < 0.05) to RIVSUM, a measure of freshwater discharge from the St Lawrence River system and an index of variability in the region's climate. Three hypotheses are put forward to explain these observations: (1) there is a strong link between interannual variation in abundance of copepod females, which produce prey for mackerel larvae, and larval survival; the exceptional recruitment and subsequent year class resulted from an exceptional production of Calanus finmarchicus nauplii; (2) years of high zooplankton biomass provide better feeding conditions and consequently higher survival of mackerel juveniles; and (3) mackerel recruitment and zooplankton biomass are independently under the control of an underlying physical process, without strong trophic linkage. The first hypothesis is supported by a study of copepod species composition and female abundance conducted for four of the survey years. At the present time, none of these hypotheses can be ruled out.     

Lipids,buoyancy and the seasonal vertical migration of Calanus finmarchicus

The copepod Calanus finmarchicus remains in diapause for up to 5 months in the cold (<0.5°C) deep (>700 m) waters of the Faroe–Shetland Channel of the north-western approaches to the North Sea. While in diapause, C. finmarchicus has a high lipid content, up to 76% of dry weight, mostly in the form of wax esters. The question we address here is how copepods with such a high content of buoyant lipids can remain in diapause at depth for an extended period of time? The corollary to this is how this lipid content hinders and/or assists the copepods in their seasonal vertical migration? Part of the answer is due to the physical properties of wax esters. These have a thermal expansion and compressibility higher than that of sea water. Thus, depending on their relative composition (i.e. wax esters/water/protein/chitin), a copepod that is positively buoyant in warm surface waters can become neutrally buoyant in cold deep water. We develop a simple three component physical model of a copepod to explore how and where they attain neutral buoyancy, how the lipid content can aid in their ascent, and what fraction of the lipids can be utilized in ascent in gonad/egg formation while maintaining observed ascent rates. As well as being an energy reserve, the results show that rather than being a barrier to vertical migration, wax esters serve as an important regulator of buoyancy.     

Distribution and abundance of overwintering Calanus finmarchicus in the Faroe–Shetland Channel

The distribution and abundance of overwintering Calanus finmarchicus in the Faroe–Shetland Channel was studied during December 1994 and January 1995. Maximum abundance of animals in the Channel was approximately 50 000 m⁻², with a peak concentration of 627 m⁻³ at a depth of 930 m. Copepodite stages IV and V accounted for > 98% of the population. A clear association was found between the horizontal and vertical distribution of animals and the Arctic water masses in the bottom of the Faroe–Shetland Channel. The Wyville–Thomson Ridge formed a barrier to the southern distribution of Arctic waters and the abundance of overwintering C. finmarchicus was 25 times lower to the south of the Ridge than to the north. Spatial variability in lipid content and composition indicated that overwintering C. finmarchicus in the southern Norwegian Sea were in poorer condition with respect to wax ester content, and in a more advanced state of emergence from overwintering, than the animals within the Channel. The overwintering stock in the Channel probably originated from the Norwegian Sea or from north of the Faroe Islands. The Faroe–Shetland Channel is an important source of animals advected into the northern North Sea in the spring (March/April). The population abundance in the Faroe–Shetland Channel was estimated to be 4.5 × 10¹⁴ individuals, which is more than adequate to account for the mean concentration of adult stages observed in the northern North Sea in April.     

The effect of vertical migration strategy on retention and dispersion in the Irish Sea during spring–summer

A biophysical model of the Irish Sea was produced to predict net horizontal movement of key taxa including small copepods (Acartia, Pseudocalanus), large copepod species (Calanus finmarchicus and C. helgolandicus), the euphausiid Meganyctiphanes norvegica, and the larval stages of commercially important fish and Nephrops. The model coupled biological information on the vertical distribution and diel vertical migration (DVM) of these taxa with temporally resolved flow fields generated by a baroclinic hydrodynamic model. The DVM of Calanus spp. and Meganyctiphanes was parameterized empirically from data provided by a biophysical campaign carried out in the nearby Clyde Sea. Small copepods and larvae remained within the layers above the thermocline. Model organisms, programmed with particular behavioural patterns, were spaced at regular intervals (1/20° longitude × 1/30° latitude) in the model grid, which extended from 53 to 54.5°N and 4.5 to 6.4°W. The model was run over the months of April, May and June. Model simulations showed that the movement of model organisms followed one of four characteristic patterns: containment along the coast, advection to the north or south, stagnation in the centre of the gyre or circulation around the periphery of the gyre. The latter pattern was apparent even in April, when gyral circulation was not expected because of a lack of stratification in the water column. Thirty‐day model runs in April, May and June showed that the number of organisms retained within the model grid varied between 23 and 49%. Those seeded close to the Irish coast had the highest probability of being retained within the model grid at the end of 90 days. DVM increased the probability of retention through improving the chances of organisms becoming entrained within the gyre circulation. This was especially true for Calanus spp., with those performing midnight sinking being the most likely to be retained out of any model organism. Meganyctiphanes showed comparatively lower levels of retention in deep regions, which suggests that they may have to swim against currents during certain phases of their DVM to avoid displacement. Overall, model results supported a number of hypotheses on the origin and fate of various taxa in the Irish Sea, as well as revealing some less expected distribution patterns.     

Lipids, buoyancy and the seasonal vertical migration of Calanus finmarchicus

The copepod Calanus finmarchicus remains in diapause for up to 5 months in the cold (<0.5°C) deep (>700 m) waters of the Faroe–Shetland Channel of the north-western approaches to the North Sea. While in diapause, C. finmarchicus has a high lipid content, up to 76% of dry weight, mostly in the form of wax esters. The question we address here is how copepods with such a high content of buoyant lipids can remain in diapause at depth for an extended period of time? The corollary to this is how this lipid content hinders and/or assists the copepods in their seasonal vertical migration? Part of the answer is due to the physical properties of wax esters. These have a thermal expansion and compressibility higher than that of sea water. Thus, depending on their relative composition (i.e. wax esters/water/protein/chitin), a copepod that is positively buoyant in warm surface waters can become neutrally buoyant in cold deep water. We develop a simple three component physical model of a copepod to explore how and where they attain neutral buoyancy, how the lipid content can aid in their ascent, and what fraction of the lipids can be utilized in ascent in gonad/egg formation while maintaining observed ascent rates. As well as being an energy reserve, the results show that rather than being a barrier to vertical migration, wax esters serve as an important regulator of buoyancy.