Barbara Niehoff

Reproduction of the Arctic copepod Calanus hyperboreus in the Greenland Sea-field and laboratory observations

The seasonal vertical migration of adultCalanus hyperboreus Krøyer and their female reproductive biology have been studied in the central Greenland Sea, based on 7-month coverage during one annual cycle. Females dwelled in surface waters only between April and July. Gonad maturation began after the summer descent into deeper waters between September and October in the absence of food. Breeding was assessed from gonad development and egg production experiments. It started around the beginning of November and was mostly completed by March/April, before the spring ascent. The overwintering depth of females ranged from close to the surface to 3000 m, and mature females had a preference for shallower depths. Males were mostly found during the breeding period, where they inhabited the 500- to 1000-m layer. Female dry weight was highest in August and November and lowest in February. Field observations were supplemented by observations on gonad development and egg production of females, which were collected in June and August and kept in the laboratory for up to 6 months. Clutch size, spawning interval and fecundity were determined at 0°C. Females collected in June were mature earlier. Due to their low dry weight, they developed only small gonads and very few of them spawned. Females from August were much fatter and produced up to 1000 eggs in several clutches deposited at regular intervals.

On birth and death in the sea

We present the first comparative study of the stage-specific patterns of mortality of Calanus and Pseudocalanus, two widely distributed genera that are representative of a relatively large-bodied, broadcast spawning calanoid copepod and a relatively small-bodied, egg-brooding calanoid. The study site is Georges Bank, a continental shelf locality in the Northwestern Atlantic with retentive circulation that renders it suitable for studies of population dynamics. Based on extensive mortality estimates from 30 cruises, we find that co-occurring Calanus finmarchicus and Pseudocalanus spp. have markedly different patterns of stage-specific mortality, the former bimodal and the latter relatively uniform with respect to developmental stage. Neither taxon exhibits a monotonic decline in mortality with developmental stage, nor are rates of mortality predictable in a useful manner by copepod body size or by ambient temperature. Young stages of the broadcast-spawning C. finmarchicus show conditional density-dependence of mortality rates, i.e. mortality rates are independent of population density when adult females are low in abundance but positively related to population density at high female abundances. This density-dependence, which is probably attributable to egg cannibalism, introduces a quadratic mortality term into population dynamic models. The egg-brooding Pseudocalanus spp., in contrast, show no evidence of density-dependent mortality. The two taxa illustrate a life history trade-off: the broadcast-spawning Calanus exhibits birth rates that are greatly elevated with respect to those of Pseudocalanus, but there is a compensatory cost in very low survivorship of the freely spawned eggs. Both the high fecundity, high mortality life history of Calanus and the low fecundity, low mortality life history of Pseudocalanus appear to have approximately equal fitness in this study site.

Oogenesis and gonad maturation in the copepodCalanus finmarchicus and the prediction of egg production from preserved samples

Oogenesis and gonad development ofCalamus finmarchicus during the winter spring transition in a fjord in Northern Norway were studied. A combination of observations on whole animals (macroscopic), taking advantage of their transparency, and histological sections was used to establish a system of gonad maturation stages, to describe the spawning cycle and to estimate egg production rates from preserved samples. During their development, oocytes change their shape and size, the morphology of the nucleus, and the appearance of the ooplasm. Four oocyte development stages were identified and related to a macroscopic system of four gonad developmental stages (GS) that can be applied to whole stained animals. During gonad development, the macroscopic morphological modifications of the gonads are mainly manifested in an increase in number and size of the oocytes and their distribution in diverticula and oviducts. The maturation processes during the spawning cycle (the period between egg depositions) were described from histological sections of females preserved at regular intervals after egg deposition. The macroscopic GS did not vary during the spawning cycle. This GS was a highly significant indicator for females, which spawned within 24 h after collection at 5°C. Another histological indicator for the last third of the spawning cycle is the appearance of distinct chromosomes, which, however, are only seen in histological sections. For the prediction of egg production rate of aCalanus population, in addition to the number of females spawning, knowledge of clutch size and the interval of clutch deposition is required. Clutch size was determined by counting the number of the oocyte stages to be released during the next spawning event in serial sections of the gonads. There was no significant difference in clutches laid by the females during 5 days before preservation. From the observations it seems that GS 4 predicts females that are ready to spawn within the minimum spawning interval characterisitic for a given temperature. If this holds true, it should be possible to predict in situ egg production rates solely from preserved samples.

The gonad morphology and maturation in Arctic Calanus species

Gonad morphology and maturation of the dominant Arctic copepod species Calanus glacialis and C. hyperboreus were studied using a combination of observations on whole animals (“whole-body analysis”) and histological sections to describe basic reproductive processes. The comparison with C. finmarchicus shows that the internal organisation of the gonads is the same in the Calanus species: from the dorsally laying ovary, two anterior diverticula extent into the head region and two posterior diverticula extend into the abdomen. In the diverticula, the oocytes increase in size and development stage in a dorso–ventral direction. There were no physiological differences between the oocytes in the diverticula and oviducts in any of the three species. During gonad development, the morphological modifications of the gonads mainly refer to an increase in number, developmental stage and size of the oocytes both in diverticula and oviducts. During spawning the most ventral layer of oocytes is released, and then the adjacent dorsal layer begins final maturation. Thus, the final maturation processes occur simultaneously as it is essential to produce clutches.

Summertime plankton ecology in Fram Strait - a compilation of long- and short-term observations

Between Greenland and Spitsbergen, Fram Strait is a region where cold ice-covered Polar Water exits the Arctic Ocean with the East Greenland Current (EGC) and warm Atlantic Water enters the Arctic Ocean with the West Spitsbergen Current (WSC). In this compilation, we present two different data sets from plankton ecological observations in Fram Strait: (1) long-term measurements of satellite-derived (1998–2012) and in situ chlorophyll a (chl a) measurements (mainly summer cruises, 1991–2012) plus protist compositions (a station in WSC, eight summer cruises, 1998–2011); and (2) short-term measurements of a multidisciplinary approach that includes traditional plankton investigations, remote sensing, zooplankton, microbiological and molecular studies, and biogeochemical analyses carried out during two expeditions in June/July in the years 2010 and 2011. Both summer satellite-derived and in situ chl a concentrations showed slight trends towards higher values in the WSC since 1998 and 1991, respectively. In contrast, no trends were visible in the EGC. The protist composition in the WSC showed differences for the summer months: a dominance of diatoms was replaced by a dominance of Phaeocystis pouchetii and other small pico- and nanoplankton species. The observed differences in eastern Fram Strait were partially due to a warm anomaly in the WSC. Although changes associated with warmer water temperatures were observed, further long-term investigations are needed to distinguish between natural variability and climate change in Fram Strait. Results of two summer studies in 2010 and 2011 revealed the variability in plankton ecology in Fram Strait.

Characteristics of digestive enzymes of calanoid copepod species from different latitudes in relation to temperature, pH and food.

In calanoid copepods it is poorly understood how enzymatic activities and patterns are affected by abiotic and biotic factors. Such knowledge, however, is crucial to assess metabolic functioning and performance of organisms in different habitats. Therefore, our study focuses on digestive enzyme activities in relation to temperature, pH and food in the Arctic species Calanus glacialis and in Centropages hamatus and Temora longicornis from the North Sea. Enzyme activities were measured over a range from 0 to 70 °C (lipases/esterases, proteinases) and pH 5 to 9 (proteinases). In all species, relative proteinases activity peaked at 40/50 °C and pH 6; relative lipases/esterases activity peaked at 30 °C. Between 0 and 20 °C, lipase activity of C. glacialis was higher (40-70% of maximum) than that of the boreal copepods (25-64%), which suggests thermal adaptation of the lipid metabolism in the polar species. Incubating C. glacialis with the diatom Thalassiosira weissflogii showed (i) that enzyme activities increased especially in the alkaline range and (ii) that enzyme patterns, revealed by gel electrophoresis, differed from that of starving individuals, indicating that feeding induced enzyme expression. Such studies, linking abiotic and biotic conditions to enzyme functioning, can help elucidating the capacity of copepods to respond to environmental changes.

Feeding by Calanus glacialis in a high arctic fjord: potential seasonal importance of alternative prey

Original Article Feeding by Calanus glacialis in a high arctic fjord: potential seasonal importance of alternative prey Alison C. Cleary*, Janne E. Søreide, Daniela Freese, Barbara Niehoff, and Tove M. Gabrielsen University Centre in Svalbard, Longyearbyen N-9171, Norway Alfred Wegener Institute, Helmholtz Centre for Polar and Marine Research, 27570 Bremerhaven, Germany *Corresponding author: tel: þ47 96 88 53 94; fax: þ47 77 75 05 01; e-mail: alison_cleary@my.uri.edu Present address: Norwegian Polar Institute, Fram Centre, Tromsø NO-9296, Norway.

Bioenergetics of the copepod Temora longicornis under different nutrient regimes

The copepod Temora longicornis depends on constant prey availability, but its performance also depends on how efficiently it utilizes its food sources. Our research goal was to understand copepod energy allocation in relation to diet quality. The working hypothesis was that Temora performs better on the diet whose elemental ratio is closest to its own. Diatoms (Diat) and dinoflagellates (Dino) cultured in nutrient-replete (+) and nitrogen-depleted (-) conditions were fed to the copepods. Ingestion, respiration, excretion and egg and fecal pellet production rates were measured. Carbon (C) and nitrogen (N) budgets were built to investigate differences in dietary C and N partitioning. Copepods fed nitrogen-depleted diatoms (Diat-), which had the most different C: N ratio to that of Temora longicornis, had high metabolic losses and low growth. Copepods fed nitrogen-rich dinoflagellates (Dino+) with a more similar C: N ratio to their own also had high metabolic losses, but displayed the highest investment into somatic growth and egg production. The results indicate that dinoflagellates are a better food source for T. longicornis. Furthermore, consumption of low-quality food leads to higher respiration rates and faster leakage of dissolved organic carbon from copepod fecal pellets; and egestion is a main pathway in copepods for eliminating unabsorbed and non-metabolized carbon.

Leben im Pelagial

Das Pelagial ist der Lebensraum des freien Wassers mit allem, was darin schwimmt und driftet, vom winzigen Bakterioplankton uber Phyto-, Zooplankton und Fische bis hin zu den den riesigen Walen. In welchen Bereichen und Tiefen leben die unterschiedlichen Organismen? Horizontale und vertikale, tages- und jahreszeitliche Wanderungen sowie die Wechselwirkungen zwischen den unterschiedlichen Organismengruppen und ihre Rolle im Nahrungsgefuge werden erlautert.