Life cycle strategies of free-living copepods in fresh waters

Freshwater copepods live in habitats characterized by a high degree of instability. To survive occasional deterioration of their environment copepods have evolved adaptive mechanisms like dormancy or migration in order to avoid lethal conditions and to synchronize growth and reproduction with favourable abiotic and biotic conditions. Typical life cycles of harpacticoid, calanoid and cyclopoid copepods are presented to show strategies that have evolved to survive threatening environmental conditions.     

Modelling the silica pump in the Permanently Open Ocean Zone of the Southern Ocean

A coupled 1D physical–biogeochemical model has been built to simulate the cycles of silicon and of nitrogen in the Indian sector of the Permanently Open Ocean Zone of the Southern Ocean. Based on a simplified trophic network, that includes two size classes of phytoplankton and of zooplankton, and a microbial loop, it has been calibrated by reference to surface physical, chemical and biological data sets collected at the KERFIX time-series station (50°40′S–68°25′E). The model correctly reproduces the high nutrient low chlorophyll features typical of the studied area. In a region where the spring–summer mixed layer depth is usually deeper than 60 m, the maximum of chlorophyll never exceeds 1.5 mg m⁻³, and the annual primary production is only 68 g C m⁻² year⁻¹. In the surface layer nitrate is never exhausted (range 27–23.5 mmoles m⁻³) while silicic acid shows strong seasonal variations (range 5–20 mmoles m⁻³). On an annual basis 71% of the primary production sustained by nanophytoplankton is grazed by microzooplankton. Compared to North Atlantic, siliceous microphytoplankton is mainly prevented from blooming because of an unfavourable spring–summer light-mixing regime. Silicic acid limitation (high half saturation constant for Si uptake: 8 mmoles m⁻³) also plays a major role on diatom growth. Mesozooplankton grazing pressure excerpts its influence especially in late spring. The model illustrates the efficiency of the silica pump in the Southern Ocean: up to 63% of the biogenic silica that has been synthetized in the photic layer is exported towards the deep ocean, while only 11% of the particulate organic nitrogen escapes recycling in the surface layer.     

Late-Quaternary changes in productivity of the Southern Ocean

Paleoceanographic records based on new proxies of export production have been constructed for the South Atlantic sector of the Southern Ocean. A radionuclide-ratio proxy of particle flux (¹⁰Be/²³⁰Th) and the accumulation rate of authigenic uranium, which responds to the flux of organic carbon to the sea bed, both indicate a dramatic increase, compared to the present, in the export production of the Subantarctic zone (approximately the region between the present-day positions of the Subtropical Convergence and the Antarctic Polar Front) during glacial periods. If the South Atlantic is representative of the entire Southern Ocean, then export production in the Southern Ocean during the Last Glacial Maximum was substantially greater than at present. Previous studies, focusing on the burial of biogenic opal, failed to recognize the glacial increase in export production of the Southern Ocean because of a strong non-linearity between accumulation rates of opal and of organic carbon.     

Primary productivity of the Palmer Long Term Ecological Research Area and the Southern Ocean

A major objective of the Palmer Long Term Ecological Research (Palmer LTER) project is to obtain a comprehensive understanding of the various components of the Antarctic marine ecosystem. Phytoplankton production plays a key role in this so-called high nutrient, low chlorophyll environment, and factors that regulate production include those that control cell growth (light, temperature, and nutrients) and those that control cell accumulation rate and hence population growth (water column stability, grazing, and sinking). Sea ice mediates several of these factors and frequently conditions the water column for a spring bloom which is characterized by a pulse of production restricted in both time and space. This study models the spatial and temporal variability of primary production within the Palmer LTER area west of the Antarctic Peninsula and discusses this production in the context of historical data for the Southern Ocean. Primary production for the Southern Ocean and the Palmer LTER area have been computed using both light-pigment production models [Smith, R.C., Bidigare, R.R., Prézelin, B.B., Baker, K.S., Brooks, J.M., 1987. Optical characterization of primary productivity across a coastal front. Mar. Biol. (96), 575–591; Bidigare, R.R., Smith, R.C., Baker, K.S., Marra, J., 1987. Oceanic primary production estimates from measurements of spectral irradiance and pigment concentrations. Global Biogeochem. Cycles (1), 171–186; Morel, A., Berthon, J.F., 1989. Surface pigments, algal biomass profiles and potential production of the euphotic layer—relationships reinvestigated in view of remote-sensing applications. Limnol. Oceanogr. (34), 1545–1562] and an ice edge production model [Nelson, D.M., Smith, W.O., 1986. Phytoplankton bloom dynamics of the western Ross Sea ice edge: II. Mesoscale cycling of nitrogen and silicon. Deep-Sea Res. (33), 1389–1412; Wilson, D.L., Smith, W.O., Nelson, D.M., 1986. Phytoplankton bloom dynamics of the Western Ross Sea ice edge: I. primary productivity and species-specific production. Deep-Sea Res., 33, 1375–1387; Smith, W.O., Nelson, D.M., 1986. Importance of ice edge phytoplankton production in the Southern Ocean. BioScience (36), 251–257]. Chlorophyll concentrations, total photosynthetically available radiation (PAR) and sea ice concentrations were derived from satellite data. These same parameters, in addition to hydrodynamic conditions, have also been determined from shipboard and Palmer Station observations during the LTER program. Model results are compared, sensitivity studies evaluated, and productivity of the Palmer LTER region is discussed in terms of its space time distribution, seasonal and interannual variability, and overall contribution to the marine ecology of the Southern Ocean.     

Size-fractionated phytoplankton biomass and species composition in the Indian sector of the Southern Ocean during austral summer

During the late austral summer of 1994, Antarctic waters were characterized by low phytoplankton biomass. Along the 62°E meridian transect, between 49°S and 67°S, chlorophyll (Chl.) a concentration in the upper 150 m was on average 0.2 mg m⁻³. However, in the Seasonal Ice Zone (SIZ) chlorophyll a concentrations were higher, with a characteristic deep chlorophyll maximum. The highest value (0.6 mg Chl. a m⁻³) was measured at the Antarctic Divergence, 64°S, corresponding to the depth of the temperature minimum (100 m). This deep biomass maximum decreased from South to North, disappeared in the Permanently Open Ocean Zone (POOZ) and reappeared with less vigour in the vicinity of the Polar Front Zone (PFZ). In the SIZ, the upper mixed layer was shallow, biomass was higher and the >10 μm fraction was predominant. In this zone the >10 μm, 2–10 μm and <2 μm size fractions represented on the average 46%, 25.1% and 28.9% of the total integrated Chl. a stock in the upper 100 m, respectively. The phytoplankton assemblage was diverse, mainly composed of large diatoms and dinoflagellate cells which contributed 42.7% and 33.1% of the autotrophic carbon biomass, respectively. Moving northwards, in parallel with the decrease in biomass, the biomass of autotrophic pico- and nanoflagellates (mainly Cryptophytes) increased steadily. In the POOZ, the picoplanktonic size fraction contributed 47.4% of the total integrated Chl. a stock. A phytoplankton community structure with low biomass and picoplankton-dominated assemblage in the POOZ contrasted with the relatively rich, diverse and diatom-dominated assemblage in the SIZ. These differences reflect the spatial and temporal variations prevailing in the Southern Ocean pelagic ecosystem.     

Life cycle of Epischura baicalensis Sars (Copepoda, Calanoida) in Lake Baikal

Epischura baicalensis Sars is a dominant pelagic species of Lake Baikal zooplankton. This is endemic to Lake Baikal and inhabits the entire water column. It produces two generations per year: the winter–spring and the summer. These copepods develop under different ecological conditions and vary in the duration of life stages, reproduction time, maturation of sex products and adult males and females lifespan. The total life period of the animals from each generation is one year. One female can produce 10 egg sacks every 10–20 days during its life time. The ratio of males and females is 1:1. One of the most essential features of the ecology of E. baicalensis is the alteration of its mass inhabited areas during a year, as well as in day time. This is due to the need for various conditions for gonad maturation, reproduction, nourishing and protection from being consumed by planktivors.     

Annual cycle of distribution of three ice-associated copepods along the coast near Dumont d'Urville, Terre Adélie (Antarctica)

In polar regions sea ice is a site of enhanced primary production during winter and provides important habitat for small grazers, such as copepods. We sampled zooplankton from the sea ice and water column throughout 2005, near Dumont d'Urville station (Terre Adélie, Antarctica). Three species of ice-associated copepods were found: two calanoid copepods Paralabidocera antarctica and Stephos longipes and the harpacticoid copepod Drescheriella glacialis. P. antarctica was the most abundant of the three and was closely associated with the sea ice during most of the year. This species had a one year life cycle with a probable over-wintering period in the sea ice as nauplii and a short copepodite phase in spring. Reproduction and spawning occurred in early summer. A comparison with two other populations (near Syowa and Davis stations) along the east coast of Antarctica showed that there was a temporal shift in the life cycles of the three populations, which was linked to variability in sea ice conditions. D. glacialis was the second most abundant copepod and was more common during the winter than during summer, indicating its preference for the sea ice habitat. In autumn, the presence of D. glacialis in the deeper part of the water column suggested that this species colonised the sea ice from the benthos. S. longipes was found only in the water column near Dumont d'Urville and was not very abundant. In Terre Adélie particular environmental conditions, such as the absence of a permanent sea ice zone throughout the year, a longer time of open water, strong katabatic winds and the presence of polynyas, have influenced both the abundance and distribution of the three common ice-associated copepods.     

Glacial–interglacial changes in the accumulation rates of major biogenic components in Southern Indian Ocean sediments

In this study we compare major biogenic components (opal-A, carbonate, and organic carbon) and authigenic uranium accumulation rates from the southeastern Indian Ocean for both Holocene and glacial periods. Integrated accumulation rates across the whole Indian sector of the Southern Ocean indicate that the burial of organic carbon which is held approximately constant, contrasts with lower biogenic silica and carbonate burial rates during glacial intervals. In addition, higher glacial accumulation rates of authigenic uranium are found in the sediments of the Polar Front Zone (PFZ) and the Sub-Antarctic zone (SAZ) than anywhere in the modern Southern Ocean. This suggests more reducing conditions in the PFZ and SAZ during the last glacial maximum. The simplest explanation of a northward shift of the PFZ cannot explain such changes. Glacial sediment burial changes result probably from deep water decrease in oxygen levels and increase in CO₂ due to combination of two processes: (1) hydrologic changes and (2) continuous organic carbon export fluxes to the seafloor. Such shifts in chemical conditions could have enhanced the dissolution of carbonates and better preserved the organic carbon in sediments, leading in significant changes of biogenic silica/C_org and CaCO₃/C_org flux ratios.     

Phytoplanktonic nutrient utilisation and nutrient signature in the Southern Ocean

The separation in Southern Ocean provinces of silicate excess at nitrate exhaustion and of nitrate excess at silicate exhaustion was already introduced by Kamykowski and Zentara (Kamykowski, D., Zentara, S.J., 1985. Nitrate and silicic acid in the world ocean: patterns and processes. Mar. Ecol. Prog. Ser. 26, 47–59; and Kamykowski, D., Zentara, S.J., 1989. Circumpolar plant nutrient covariation in the Southern Ocean: patterns and processes. Mar. Ecol. Prog. Ser. 58, 101–111) and our investigations of the silicate to nitrate uptake ratios confirm the earlier distinction. Oligotrophic antarctic waters mainly exhibit proportionally higher silicate removal what induces a potential for nitrate excess. The nitrogen uptake regime of such areas is characterised by low absolute as well as specific nitrate uptake rates throughout. Maximal values did not exceed 0.15 μM d⁻¹ and 0.005 h⁻¹, respectively. Corresponding f-ratios ranged from 0.39 to 0.86. This scenario contrasts strikingly to the more fertile ice edge areas. They showed a drastic but short vernal increase in nitrate uptake. Absolute uptake rates reached a maximum value of 2.18 μM d⁻¹ whereas the maximal specific uptake rate was 0.063 h⁻¹. In addition to an optimal physical environment for bloom development, accumulation of ammonium stimulated nitrate uptake in a direct or indirect way. Since ammonium build-up in surface waters traces enhanced remineralisation, release of other essential compounds during degradation of organic matter might have been the main trigger. This peak nitrate utilisation during early spring led to the observed potential for silicate excess. With increasing seasonal maturity the nitrate uptake became inhibited by the presence of enhanced ammonium availability (up to 8% of the inorganic nitrogen pool), however, and after a short period of intensive nitrate consumption the uptake rates drop to very low levels, which are comparable to the ones observed in the area of nitrate excess at silicate exhaustion.     

The role of copepods in the Baikal ecosystem

This describes a study of the pelagic zone of Lake Baikal with particular reference to copepods. In addition the cycling of matter and energy in lacustrine basins is described.     

Patterns in diversity and community structure of epipelagic copepods from the Brazil–Malvinas Confluence area, south-western Atlantic

Diversity and structure of epipelagic copepod communities were investigated using 70 zooplankton samples collected from the top 50 m of the Brazil–Malvinas Confluence area between 35° and 55°S. Biogeographic patterns were investigated using multivariate analysis. Biodiversity patterns were examined using different univariate indexes. Representatives of 35 species of copepods from 23 genera and 13 families were found. Two zones were delimited based on their copepod composition, one in the subtropical waters of the Brazil current, and the other in the subantarctic waters of the Malvinas current. Both environments displayed contrasting patterns of biodiversity depending upon which element is measured (regional species richness as well as mean point species richness were significantly higher in the subtropical group of stations, whereas taxonomic distinctness was significantly higher in the subantarctic community). Based on these contrasting results, we suggest the use of both kind of indexes when defining priority areas for conservation.     

On the relation of structure, perception and activity in marine planktonic copepods

The goal of this paper is to illustrate how in juvenile and adult subtropical marine planktonic copepods various structures or morphological features function in concert to detect prey and predators. Without motion by either food (e.g. flagellate, ciliate) or feeder (e.g. feeding current) or both (e.g. Acartia spp. and ciliate) few feeding activities will occur. Through motion a food particle is either perceived mechanically or chemically to be followed by appendage activities. A combination of mechano- and chemosensors on their cephalic appendages (and probably on other extremities) serve juvenile and adult copepods to perceive signals. Perception is followed by alternation of motion and sensing by these appendages, or by no motion at all (e.g. behavior by Eucalanus pileatus when perceiving a weak hydrodynamic signal). Non-moving and extended sensors (setae) are best suited for mechanical/hydrodynamic perceptions in those copepods which lack a feeding current and hardly move. Numerous mechanosensors arranged in three dimensions on the first antennae (A1) are required to perceive the precise location of moving prey at a distance (e.g. Oithona feeding on ciliates but also sinking particles). Those copepods which create a weak or intermittent feeding current can supplement nutrition with carnivory, which requires perception by the A1 (e.g. Centropages velificatus adults). These two groups require, in addition to perception of prey motion/location, rapid motion by their appendages (A1, second maxillae M2, etc.) to capture the prey. Nauplii, which satiate at far lower food levels than adults, have one of several means of food acquisition: encounter through forward motion, perception through feeding current, or perception of a moving food particle. The nearly continuous motion of most calanoid nauplii makes them vulnerable to predation because all three pairs of appendages are usually moving. Opposite are nauplii of cyclopoid and a few calanoid species which move only occasionally. Copepodid stages and adults use non-moving and often extended setae on the tips of their A1 to perceive predators at a distance. This structure and their pronounced escape motion may reduce their vulnerability to predation as compared to nauplii.     

Ichthyoplankton in the neritic and coastal zone of Antarctica and Subantarctic islands: A review

Since the article published by Loeb et al. [Loeb, V.J., Kellermann, A., Koubbi, P., North, A.W., White, M., 1993. Antarctic larval fish assemblages: a review. Bull. Mar. Sci. 53(2), 416–449.] about Antarctic ichthyoplankton, many surveys were carried out in different sectors of the Southern Ocean focusing on different aspects of the ecology of fish larvae. Some of these researches were conducted in the Subantarctic Kerguelen Islands and others on the continental shelf off Terre Adélie and Georges V land. Oceanographic and geographic features influence fish larvae ecology such as island mass effects, gyres, canyons. Antarctic fishes show also temporal segregation of spawning which induces temporal succession of early stage larvae. This avoids competition and probably the predation on early stages for species having few recruits. In that case, we have to understand how these larvae can deal with the match–mismatch with their preys and how they find sufficient food to survive. But our knowledge on Antarctic fish larvae is still insufficient as we do not know larvae for quite a lot of species and because of the difficulty to sample during winter.     

Community structure of mesopelagic copepods (>500 μm) in the Ligurian Sea (Western Mediterranean)

The distribution of mesopelagic copepods between 250 and 1000 m depth was studied along a transect off the coast of Villefranche-sur-Mer in June 1991 by means of a BIONESS multiple-net sampler. Among the most abundant species at all stations were Pleuromamma gracilis, Paraeuchaeta acuta and overwintering Calanus helgolandicus CV copepodites, each species inhabiting different depth layers with maximum abundances in the 250–350 m, 450–550 m and the 700–850 m layer, respectively. Cluster analysis of the vertical distribution of all species caught revealed three distinct strata (‘TOP', ‘INTERMEDIATE', ‘BOTTOM') consisting of characteristic species that occurred in the same stratum at all stations. Differences in total abundances of the species assemblage occupying each layer are discussed with regard to predation pressure by mesopelagic macroplanktonic crustaceans and fish. We show that each stratum is characterized by distinct trophic interactions and life strategies, which play an important role in the structuring of mesopelagic zooplankton communities.     

Fluxes of biogenic carbon in the Southern Ocean: roles of large microphagous zooplankton

The Southern Ocean is an extreme environment, where waters are permanently cold, a seasonal ice cover extends over large areas, and the solar energy available for photosynthesis is severely restricted, either by vertical mixing to considerable depths or, especially south of the Antarctic Circle, by prolonged seasonal periods of low or no irradiance. Such conditions would normally lead to low productivity and a water column dominated by recycling processes involving microbial components of pelagic communities but this does not seem to be the case in the Southern Ocean, where there is efficient export to large apex predators and deep waters. This paper investigates the role of large microphagous zooplankton (salps, krill, and some large copepods) in the partitioning of biogenic carbon among the pools of short- and long-lived organic carbon and sequestered biogenic carbon. Large microphagous zooplankton are able to ingest microbial-sized particles and thus repackage small, non-sinking particles into both metazoan biomass and large, rapidly sinking faeces. Given the wide spatio-temporal extent of microbial trophic pathways in the Southern Ocean, large zooplankton that are omnivorous or able to ingest small food particles have a competitive advantage over herbivorous zooplankton. Krill efficiently transfer carbon to a wide array of apex predators and their faecal pellets are exported to depth during occasional brief sedimentation episodes in spring time. Salps may be a significant link towards some fish (directly) and other apex predators (indirectly) and, at some locations (especially in offshore waters) and time, they may account for most of the downward flux of biogenic carbon. Large copepods are a trophic link towards fish and at least one whale species, and their grazing activity generally impedes the export of organic particles to depth. As a result, biogenic carbon is channelled mainly towards apex predators and episodically into the deep ocean. Without these original interactions, Antarctic waters might well be dominated by microbial components and recycling processes instead of active export from the generally small primary producers towards large apex predators.     

Study of the seasonal cycle of the biogeochemical processes in the Ligurian Sea using a 1D interdisciplinary model

A one-dimensional coupled physical–biogeochemical model has been built to study the pelagic food web of the Ligurian Sea (NW Mediterranean Sea). The physical model is the turbulent closure model (version 1D) developed at the GeoHydrodynamics and Environmental Laboratory (GHER) of the University of Liège. The ecosystem model contains 19 state variables describing the carbon and nitrogen cycles of the pelagic food web. Phytoplankton and zooplankton are both divided in three size-based compartments and the model includes an explicit representation of the microbial loop including bacteria, dissolved organic matter, nano-, and microzooplankton. The internal carbon/nitrogen ratio is assumed variable for phytoplankton and detritus, and constant for zooplankton and bacteria. Silicate is considered as a potential limiting nutrient of phytoplankton's growth. The aggregation model described by Kriest and Evans in (Proc. Ind. Acad. Sci., Earth Planet. Sci. 109 (4) (2000) 453) is used to evaluate the sinking rate of particulate detritus. The model is forced at the air–sea interface by meteorological data coming from the “Côte d'Azur” Meteorological Buoy. The dynamics of atmospheric fluxes in the Mediterranean Sea (DYFAMED) time-series data obtained during the year 2000 are used to calibrate and validate the biological model. The comparison of model results within in situ DYFAMED data shows that although some processes are not represented by the model, such as horizontal and vertical advections, model results are overall in agreement with observations and differences observed can be explained with environmental conditions.     

Patterns and controls of chlorophyll-a and primary productivity cycles in the Southern California Bight

Factors that influence the magnitude and the depth of the chlorophyll maximum layer in the ocean off Southern California are explored using observations from the long-term California Cooperative Oceanic Fisheries Investigations (CalCOFI) program. The data record is sufficiently long to reveal patterns not always evident from single stations or single cruises. Processes such as coastal wind-driven upwelling, geostrophic circulation, and annual physical and chemical cycles are illustrated to demonstrate their effect on euphotic zone nutrient availability, and subsequent phytoplankton biomass and primary productivity. In this area, where the influence of wind-driven upwelling is spatially restricted and advected waters are generally nutrient depleted, geostrophically induced upwelling and winter convection become important in determining spatial and temporal patterns of phytoplankton.     

The trophic role of the tunicate Salpa thompsoni in the Antarctic marine ecosystem

During a repeat grid survey and drogue study carried out in austral summer 1994/95, the abundance and feeding activity of salps were estimated in the Lazarev Sea region from net tows and in situ measurements of gut fluorescence. Throughout the survey area, Salpa thompsoni accounted for >95% of the total salp stock while Ihlea racovitzai was consistently represented in very low abundances. Maximum densities of S. thompsoni, with ≈4000 ind. 1000 m⁻³, were recorded in the Marginal Ice Zone (MIZ) in December when chlorophyll-a concentrations were well below 1 mg m⁻³. A dramatic decrease in salp stock was observed at the beginning of January, when S. thompsoni virtually disappeared from the most productive area of the MIZ where chlorophyll-a concentrations had by then reached bloom levels of 1.5–3 mg (Chl-a) m⁻³. In situ grazing measurements showed that throughout the cruise S. thompsoni exhibited the highest ingestion rates per individual of any of the most abundant components of the grazing pelagic community, with maxima of ≈160 μg (pigm) ind. ⁻¹ d⁻¹. These feeding rates are 3 to 5 times higher than those previously obtained using in vitro incubations. The total daily consumption of the population of S. thompsoni varied from 0.3 to 108% of daily primary production. We suggest that competitive removal of food by S. thompsoni, rather than direct predation, is responsible for the low krill abundances generally associated with salp swarms.     

Determination of anthropogenic CO2 in the North Atlantic Ocean using water mass ages and CO2 equilibrium chemistry

A new method to calculate the anthropogenic CO₂ (ΔDIC_ant) within the water column of the North Atlantic Ocean is presented. The method exploits the equilibrium chemistry of the carbonate system with reference to temperature, salinity and the partial pressure of atmospheric CO₂ (pCO_2,atm). ΔDIC_ant is calculated with reference to the ventilation ages of water masses derived from tracer data and to the time history of pCO_2,atm. The method is applied to data recorded during the WOCE program on the WHP A1/E transect in the North Atlantic Ocean, where we characterise six key water masses by their relationships of dissolved inorganic carbon (DIC) and apparent oxygen utilisation (AOU). The error in determining ΔDIC_ant is reduced significantly by minimising the number of values referred to, especially by avoiding any use of remineralisation ratios of particulate organic matter. The distribution of ΔDIC_ant shows highest values of up to 45 μmol kg⁻¹ in the surface waters falling to 28–33 μmol kg⁻¹ in the Irminger Sea west of the Mid-Atlantic Ridge. The eastern basin is imprinted by older water masses revealing decreasing values down to 10 μmol kg⁻¹ ΔDIC_ant in the Antarctic Bottom Water. These findings indicate the penetration of the whole water column of the North Atlantic Ocean by anthropogenic CO₂.     

Carbonate chemistry dynamics and carbon dioxide fluxes across the atmosphere–ice–water interfaces in the Arctic Ocean: Pacific sector of the Arctic

Climatic changes in the Northern Hemisphere have led to remarkable environmental changes in the Arctic Ocean, which is surrounded by permafrost. These changes include significant shrinking of sea-ice cover in summer, increased time between sea-ice break-up and freeze-up, and Arctic surface water freshening and warming associated with melting sea-ice, thawing permafrost, and increased runoff. These changes are commonly attributed to the greenhouse effect resulting from increased atmospheric carbon dioxide (CO₂) concentration and other non-CO₂ radiatively active gases (methane, nitrous oxide). The greenhouse effect should be most pronounced in the Arctic where the largest air CO₂ concentrations and winter–summer variations in the world for a clean background environment were detected. However, the air–land–shelf interaction in the Arctic has a substantial impact on the composition of the overlying atmosphere; as the permafrost thaws, a significant amount of old terrestrial carbon becomes available for biogeochemical cycling and oxidation to CO₂. The Arctic Ocean's role in determining regional CO₂ balance has been ignored, because of its small size (only  4% of the world ocean area) and because its continuous sea-ice cover is considered to impede gaseous exchange with the atmosphere so efficiently that no global climate models include CO₂ exchange over sea-ice. In this paper we show that: (1) the Arctic shelf seas (the Laptev and East-Siberian seas) may become a strong source of atmospheric CO₂ because of oxidation of bio-available eroded terrestrial carbon and river transport; (2) the Chukchi Sea shelf exhibits the strong uptake of atmospheric CO₂; (3) the sea-ice melt ponds and open brine channels form an important spring/summer air CO₂ sink that also must be included in any Arctic regional CO₂ budget. Both the direction and amount of CO₂ transfer between air and sea during open water season may be different from transfer during freezing and thawing, or during winter when CO₂ accumulates beneath Arctic sea-ice; (4) direct measurements beneath the sea ice gave two initial results. First, a drastic pCO₂ decrease from 410 μatm to 288 μatm, which was recorded in February–March beneath the fast ice near Barrow using the SAMI-CO₂ sensor, may reflect increased photosynthetic activity beneath sea-ice just after polar sunrise. Second, new measurements made in summer 2005 beneath the sea ice in the Central Basin show relatively high values of pCO₂ ranging between 425 μatm and 475 μatm, values, which are larger than the mean atmospheric value in the Arctic in summertime. The sources of those high values are supposed to be: high rates of bacterial respiration, import of the Upper Halocline Water (UHW) from the Chukchi Sea (CS) where values of pCO₂ range between 400 and 600 μatm, a contribution from the Lena river plume, or any combination of these sources.