Hans-Jürgen Hirche

Diapause in the marine copepod, Calanus finmarchicus — A review

Abstract Overwintering strategy plays a key role in the life cycle of Calanus Jinmarchicus. This species has developed adaptations such as arrested development, ontogenetic vertical migration and reduced metabolism in late developmental stages in order to survive long periods of food shortage, typical of temperate and high latitudes. At the same time preparations for the reproductive period are an integral part of its overwintering physiology. Terms such as resting phase or stage, dormancy, quiescence, hibernation and diapause are frequently encountered in descriptions of the overwintering conditions, but have not been defined. As many aspects of insect diapause are also found in the overwintering physiology of C. Jinmarchicus, an attempt was made to classify patterns of overwintering physiology according to insect diapause phases. A short comparison was also made with the diapause of freshwater cyclopoids. Although, with the knowledge available, it is difficult in some cases to correlate physiological pa...

Density-dependent mortality in an oceanic copepod population.

Planktonic copepods are primary consumers in the ocean and are perhaps the most numerous metazoans on earth. Secondary production by these zooplankton supports most food webs of the open sea, directly affecting pelagic fish populations and the biological pump of carbon into the deep ocean. Models of marine ecosystems are quite sensitive to the formulation of the term for zooplankton mortality, although there are few data available to constrain mortality rates in such models. Here we present the first evidence for nonlinear, density-dependent mortality rates of open-ocean zooplankton. A high-frequency time series reveals that per capita mortality rates of eggs of Calanus finmarchicus Gunnerus are a function of the abundance of adult females and juveniles. The temporal dynamics of zooplankton populations can be influenced as much by time-dependent mortality rates as by variations in ‘bottom up’ forcing. The functional form and rates chosen for zooplankton mortality in ecosystem models can alter the balance of pelagic ecosystems, modify elemental fluxes into the oceans interior, and modulate interannual variability in pelagic ecosystems.

Egg production and lipid content of Calanus glacialis in spring: indication of a food-dependent and food-independent reproductive mode

Female Calanus glacialis were collected in early May 1989 in the pack ice region of the western Barents Sea and were fed or starved over 11 wk. Both groups laid eggs continuously during this period, however, fed females laid up to six times more eggs. During the first 10 d after collection, both groups spawned at low rates. There-after, fed females strongly increased spawning rates and maintained high egg production levels over 11 wk, while the rates of starved females decreased. During starvation they lost 70% body carbon, 50% body nitrogen and 70% lipids. The wax ester portion decreased from 86 to ca. 60% of total lipids. Three phases of gonad development and lipid metabolism were distinguished: early gonad development; gonad maturation with a rapid decrease in lipids, especially wax esters; and spawning under fed and starved conditions, where in fed females food provided most of the energy, whereas in starved females the lipid content strongly decreased.

Distribution of dominant copepods in the Nansen Basin, Arctic Ocean, in summer

Zooplankton were collected from five depth strata (0-25-50-100-200-500 m) at 12 stations along two sections across the Nansen Basin of the Arctic Ocean in July/August 1987. Vertical and horizontal distribution of biomass and abundance of the four dominant copepod species Calanus finmarchicus, C. glacialis, C. hyperboreus and Metridia longa, together with their gonad maturity state, are presented. Egg production of C. finmarchicus, C. glacialis and Metridia longa was measured from concurrent Bongo net samples in the upper 80 m. Four biological provinces were distinguished, which closely agreed with the hydrographic regimes. In the south, the Barents Shelf slope and the Southern Transition were located in the boundary current system of Atlantic water flowing eastward along the Eurasian shelves. They were characterized by the dominance of Calanus finmarchicus. The Barents Shelf slope, representing the core of the boundary current, hosted large populations of C. hyperboreus and Metridia longa. Metridia longa, with its main occurrence in the depth of the Atlantic layer, was the biological marker of this province. The southern provinces were separated from the Central Nansen Basin and the Nansen-Gakkel Ridge by a sharp discontinuity in biomass, stage distribution and gonad maturity of the three Calanus species north of 83°N. This faunistic boundary coincided with a well defined hydrographic frontal zone. Apparently, zooplankton distribution followed closely the flow of Atlantic water below 60-200 m eastward, although the three Calanus species had their center of abundance in the surface layer consisting of polar water, which supposedly follows the Transpolar Drift westward.

Temperature impact on reproduction and development of congener copepod populations

Abstract The goal of this study was to relate the temperature response of all developmental stages and reproductive biology of two congener copepod pairs inhabiting different biogeographic regions to their geographic distribution patterns. Survival of adult females and egg production, embryonic development and hatching success of the genera Centropages and Temora from two stations, in the North Sea and the Mediterranean, were studied in laboratory experiments in a temperature range from 2 to 35 °C. Postembryonic development was determined from cohorts raised at temperatures between 10 and 20 °C with surplus food. Tolerance limits and optima of female survival, reproduction and development distinguished the northern species Centropages hamatus and Temora longicornis from the southern T. stylifera, while C. typicus, which is found in both regions, was intermediate. Thus, thermal preferences could in part explain distribution patterns of these species. While C. hamatus and the two Temora species showed distinct temperature ranges, C. typicus was able to tolerate different temperature conditions, resulting in its wide distribution range from the subarctic to the tropics. However, the thermal range of a species did not necessarily correlate with the optimal temperatures in the experiments. Optima of egg production and stage development were surprisingly low in T. stylifera, which has a mere southern distribution.

The reproductive biology of the marine copepod, Calanus finmarchicus — A review

Abstract This review of the reproductive biology of Calanus finmarchicus includes a description of the reproductive system and its development and biochemical aspects of gonadogenesis and oogenesis. Spawning and the factors controlling it, aspects of the timing of reproduction, and applications of egg production measurement are also dealt with.

Plankton distribution and the impact of copepod grazing on primary production in Fram Strait, Greenland Sea

Two hydrobiological transects across the East Greenland Shelf and the open waters of Fram Strait in summer were chosen to illustrate the distribution and production of phyto- and zooplankton in relation to water masses and ice cover. The parameters used were temperature and salinity, inorganic nutrients, chlorophyll a, primary production, phytoplankton species composition, abundance of the dominant herbivorous copepods Calanus finmarchicus, C. glacialis, C. hyperboreus and Metridia longa, and egg production of C. finmarchicus and C. glacialis. Grazing impact of copepodites and adults of these four species was modelled for each station by using egg production rates as an index of growth. Seasonal development of plankton communities was closely associated with the extent of the ice cover, hydrographic conditions and the water masses typical of the different hydrographic domains. Four regions were identified from their biological acivities and physical environment: The Northeast Water polynya on the East Greenland Shelf, with a springbloom of diatoms and active reproduction of herbivorous copepods. The pack ice region, dominated by small flagellates and negligible grazing activities. The marginal ice zone, with high variability and strong gradients of autotroph production related to eddies and ice tongues, an active microbial loop and low egg production. The open water, with high station-to-station variability of most of the parameters, probably related to hydrographic mesoscale activities. Here, Phaeocystis pouchetii was a prominent species in the phytoplankton communities. Its presence may at least partly be responsible for the generally low egg production in the open waters. Grazing impact on primary production was always small, due to low zooplankton biomass in the polynya, and due to low ingestion in the remaining regions.

Distribution, reproduction and development of Calanus species in the Northeast water in relation to environmental conditions

Abstract The vertical and horizontal distribution of mesozooplankton biomass and its composition, together with the reproduction and development of the three dominant herbivorous copepods Calanus finmarchicus, C. glacialis and C. hyperboreus , were studied in the Northeast Water between the end of May and end of July, 1993. Biomass varied between 0.15 and 10.6 g m −2 and was lower on the shallow banks. Highest biomass was found on the shelf slope and in the entrances to Belgica and Westwind Troughs. There was no seasonal trend during the study period. Among the zooplankton taxa, copepods were dominant, with 84% biomass of all other taxa, followed by chaetognaths with 14%. The large Calanus species made up 91% of copepod biomass. The most abundant species, C. glacialis , inhabited areas of low current speeds on Belgica and Ob Bank, C. hyperboreus dominated shelf slope and trough stations, while C. finmarchicus was most abundant in the Return Atlantic Current along the shelf slope and also eastern Belgica Trough. C. glacialis was the only Calanus species spawning during this period, but young copepodites of the other species were also abundant. Egg production of C. glacialis was at a maximum by our arrival and continued to at least mid August. According to the results from starvation experiments, its egg production was fuelled by food uptake, but was decoupled from phytoplankton chlorophyll until July, indicating ice-algae and microzooplankton as an alternative food source. Only when the polynya approached its maximum extent was a close relationship to phytoplankton established. Due to both spatial and temporal heterogeneity of the occurrence of young copepodite stages it was impossible to follow the growth of cohorts of developmental stages. Herbivorous carbon requirements estimated from egg production rates were mostly less than one third of the phytoplankton stock. From egg production and the distribution of young stages, the outer Westwing Trough seems to be the centre of biological activities. This may be related to the formation of young ice in winter in this area, which was found to carry a large mass of under-ice algae. High biomass but low production in Belgica Trough indicate this as an advective and expatriated community of C. finmarchicus and C. hyperboreus , where grazing is negligible, at least during early summer.

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.

Egg production ofCalanus finmarchicus at low temperature

Reproduction ofCalanus finmarchicus Gunnerus collected in June 1988 in Polar water and in April 1989 in Atlantic water was studied. Single females were kept at 0°C in the laboratory for 22 d (Polar) and 77 d (Atlantic) with superabundant food concentration (> 400µg Cl−1) of the diatomThalassiosira antarctica. There was no significant difference between the two populations, although more spent females were found in Polar water, probably due to the different dates of collection. The hypothesis of low temperature determining the geographic range ofC. finmarchicus via reproductive failure is not supported. Mean daily egg production rate of all females from Atlantic water over a 60 d period was 24.4, corresponding to 5.5% body C female−1 d−1, when an egg carbon content of 0.23µg is assumed. Coefficient of variation was 25%. Maximum values were 53.2 eggs female−1 d−1, corresponding to 12.1% body C d−1. The highest number of eggs spawned by a single female was 3101, corresponding to a seven-fold turnover of body C during the investigation period; >20% of females produced > 2000 eggs. Body carbon content did not change significantly during the experiment; the C:N ratio increased slightly, indicating lipid accumulation. Delay of response to starvation periods of 2, 4 and 7 d duration was always 2 d: egg production ceased 2 d after the onset of starvation and continued 2 d after onset of feeding.