Heterotrophic bacterial production in the Cretan Sea (NE Mediterranean)

In March and September 1995, bacterial production was measured by the ³H-leucine method in the oligotrophic Cretan Sea (Aegean Sea, Eastern Mediterranean) in the framework of the CINCS/MTP program. Samples were obtained from four stations (a coastal, a continental shelf and 2 open-sea stations) for the construction of vertical profiles of bacterial abundance and production. Bacterial production ranged from 0.1 μg C m⁻³ h⁻¹ at 1500 m depth, to 82 μg C m⁻³ h⁻¹ in March at 50 m at the coastal station. Higher bacterial integrated production was observed in March at the coastal station (131 mg C m⁻² d⁻¹ for the 0–100 m layer). Bacterial production, integrated through the water-column, was similar in March and September for the open-sea stations (60–70 mg C m⁻² d⁻¹). Relative to production, bacterial concentrations varied little between stations and seasons ranging from 9×10⁵ ml⁻¹ to 3×10⁵ ml⁻¹. Relationships between bacterial biomass and bacterial production indicated seasonal differences, likely reflecting resource limitation of bacterial biomass in March (bloom situation), and predator limitation of bacterial biomass in September (post-bloom situation).     

Influence of upwelling and river runoff interaction on phytoplankton assemblages in a Middle Galician Ria and Comparison with northern and southern rias (NW Iberian Peninsula)

The first oceanographic research (hydrography, nutrient salts, chlorophyll, primary production and phytoplankton assemblages) in a Middle Galician Ria was carried out in Corme-Laxe during 2001, just a year before the Prestige oil spill, being the only reference to evaluate eventual changes in the phytoplankton community. Due to the small size of this ria (6.5 km²), oceanographic processes were driven by the continental water supplied by Anllons River during the wet season (20–30 m³ s⁻¹ in winter), and the strong oceanic influence from the nearby shelf during the dry season. The annual cycle showed a spring bloom with high levels of chlorophyll (up to 14 μg Chl-a L⁻¹) and primary production (3 g C m⁻² d⁻¹) and a summer upwelling bloom (up to 8 μg Chl-a L⁻¹ and 10 g C m⁻² d⁻¹) where the proximity of the Galician upwelling core (<13.5 °C at sea surface) favors the input of upwelled seawater (up to 9 μM of nitrate and silicate) to the bottom ria layer, even during summer stratification events (primary production around 2 g C m⁻² d⁻¹). Thus, phytoplankton assemblages form a “continuum” from spring to autumn with a predominance of diatoms and overlapping species between consecutive periods; only in autumn dinoflagellates and flagellates characterized the phytoplankton community. In the Middle Rias as Corme-Laxe, the nutrient values, Chl-a, primary production and phytoplankton abundance for productive periods were higher than those reported for the Northern (Ria of A Coruña) and Southern Rias (Ria of Arousa) for year 2001; this suggests the importance of the hydrographic events occurring in the zone of maximum upwelling intensity of the Western Iberian Shelf, where a lack of annual cycles studies exists.     

Functional roles of interzonal migrating mesozooplankton in the western subarctic Pacific

Grazing experiments and production estimation based on life-history analysis of Neocalanus copepods (N. cristatus, N. plumchrus and N. flemingeri) were carried out in the Oyashio region to understand the carbon flows associated with the interzonal migrating copepods. These copepods, and also Eucalanus bungii, fed on nano- and micro-sized organisms non-selectively throughout the season. However, diatoms were the dominant food resource until May and organisms, such as ciliates were the major resource after May. Daily growth rate was estimated from the Ikeda–Motoda, Huntley–Lopez and Hirst–Sheader models. Since the growth rates were considered to be overestimates for the Huntley–Lopez model and underestimates for the other two models, we applied the weight-specific growth rates previously reported for these species in the Bering Shelf. Surface biomass of Neocalanus increased rapidly in June during the appearance of C5, and a successive increase of overwintering stock was evident in the deeper layer. The deep biomass decreased gradually from September to May during the dormant and reproduction period. N. cristatus has the largest annual mean biomass (2.3 gC m⁻²), followed by N. plumchrus (1.1) and N. flemingeri (0.4). Daily production rate of Neocalanus varied from 0.4 to 363.4 mgC m⁻² day⁻¹, to which N. cristatus was the largest contributor. Annual production was estimated as 11.5 gC m⁻² year⁻¹ for N. cristatus, 5.7 for N. plumchrus and 2.1 for N. flemingeri, yielding annual P/B ratio of 5 for each species. The annual production of Neocalanus accounted for 13.2% of the primary production in the Oyashio region. Their fecal pellets were estimated to account for 14.9% (0.7 gC m⁻² year⁻¹) of sinking flux of organic carbon at 1000-m depth. Moreover, their export flux by ontogenetic vertical migration, which is not measured by sediment trap observations, is estimated to be 91.5% (4.3 gC m⁻² year⁻¹) of carbon flux of sinking particles at 1000-m depth. These results suggest the important role of interzonal migrating copepods in the export flux of carbon.     

The European coastal zone: characterization and first assessment of ecosystem metabolism

The geomorphic, oceanographic, terrestrial and anthropogenic attributes of the European coastal zone are described and published data on ecosystem function (primary production and respiration) are reviewed. Four regions are considered: the Baltic Sea, Mediterranean Sea, Black Sea and the European Atlantic coast including the North Sea. The metabolic database (194 papers) suffers from a non-homogeneous geographical coverage with no usable data for the Black Sea which was therefore excluded from this part of our study. Pelagic gross primary production in European open shelves is, by far, the most documented parameter with an estimated mean of 41 mmol C m⁻² d⁻¹, the lowest value is reported in the Mediterranean Sea (21 mmol C m⁻² d⁻¹) and the highest one in the Atlantic/North Sea area (51 mmol C m⁻² d⁻¹). Microphytobenthic primary production, mostly measured in shallow areas, is extrapolated to the entire 0–200 m depth range. Its contribution to total primary production is low in all regions (mean: 1.5 mmol C m⁻² d⁻¹). Although macrophyte beds are very productive, a regional production estimate is not provided in this study because their geographical distribution along the European coastline remains unknown. Measurements of pelagic community respiration are clearly too sparse, especially below the euphotic zone, to yield an accurate picture of the fate of organic matter produced in the water column. With a mean value of 17 mmol C m⁻² d⁻¹, benthic community respiration consumes approximately 40% of the pelagic organic matter production. Estuaries generally exhibit high metabolic rates and a large range of variation in all parameters, except microphytobenthic primary production. Finally, the problem of eutrophication in Europe is discussed and the metabolic data obtained in the framework of the Land–Ocean Interactions in the Coastal Zone (LOICZ) project are compared with available direct measurements of net ecosystem production.     

Macrofaunal density and biomass in the Campeche Canyon, Southwestern Gulf of Mexico

The composition, density and community structure of the benthic macrofauna were investigated in sediments of the Campeche Canyon in the SW Gulf of Mexico. Total macrofaunal density ranged from 9466±2736 ind m⁻² at the continental shelf station to 1550±195 ind m⁻² in the canyon. Density values significantly diminished with distance from the coast and depth; only a few stations in the center of the canyon displayed larger density values (E-37 with 4666±1530 ind m⁻², E-36 with 5791±642 ind m⁻² and E-26 with 6925±2258 ind m⁻²). Densities were positively correlated to organic nitrogen in the sediment (r=0.82) and coarse silt (r=0.43), and negatively with depth (r=−0.74) and distance from the coast (r=−0.68). At all stations, the polychaete worms contributed most to the multi-species community structure. The nematodes and Foraminifera displayed their highest densities in the center of the canyon. The biomass values declined significantly with depth. We conclude that the macrofauna density and biomass changed in response to organic matter contents in the sediment, both with distance from the coast and with depth.     

Changes in sediment processes across the western Irish Sea front

Sediment characteristics, sediment respiration (oxygen uptake and sulphate reduction) and sediment–water nutrient exchange, in conjunction with water column structure and phytoplankton biomass were measured at five stations across the western Irish Sea front in August 2000. The transition from thermally stratified (surface to bottom temperature difference of 2.3 °C) to isothermal water (14.3 °C) occurred over a distance of 13 km. The influence of the front on phytoplankton biomass was limited to a small region of elevated near surface chlorophyll (2.23 mg m⁻³; 50% > biomass in mixed waters). The front clearly marked the boundary between depositional sediments (silt/clays) with elevated sediment pigment levels (≈60 mg m⁻²) on the western side, to pigment impoverished (<5 mg m⁻²) sand, through to coarse sand and shell fragments on the eastern side. Maximal rates of sedimentary respiration on the western stratified side of the front e.g. oxygen uptake S2 (852 μmol O₂ m⁻² h⁻¹) and sulphate reduction at S1 (149 μmol SO₄²⁻ m⁻² h⁻¹), coupled to significant efflux of nitrate and silicate at the western stations indicate closer benthic–pelagic coupling in the western Irish Sea. Whether this simply reflects the input of phytodetritus from the overlying water column or entrapment and settlement of pelagic production from other regions of the Irish Sea cannot yet be resolved.     

Mesozooplankton biomass and grazing responses to Cyclone Opal, a subtropical mesoscale eddy

As part of E-Flux III cruise studies in March 2005, plankton net collections were made to assess the effects of a cyclonic cold-core eddy (Cyclone Opal) on the biomass and grazing of mesozooplankton. Mesozooplankton biomass in the central region of Cyclone Opal, an area of uplifted nutricline and a subsurface diatom bloom, averaged 0.80±0.24 and 1.51±0.59 g DW m⁻², for day and night tows, respectively. These biomass estimates were about 80% higher than control (OUT) stations, with increases more or less proportionately distributed among size classes from 0.2 to >5 mm. Though elevated relative to surrounding waters south of the Hawaiian Islands (Hawai’i lee), total biomass and size distribution in Cyclone Opal were almost exactly the same as contemporary measurements made at Stn. ALOHA, 100 km north of the islands, by the HOT (Hawaii Ocean Time-series) Program. Mesozooplankton biomass and community composition at the OUT stations were also similar to ALOHA values from 1994 to 1996, preceding a recent decadal increase. These comparisons may therefore provide insight into production characteristics or biomass gradients associated with decadal changes at Stn. ALOHA. Gut fluorescence estimates were higher in Opal than in ambient waters, translating to grazing impacts of 0.11±0.02 d⁻¹ (IN) versus 0.03±0.01 d⁻¹ (OUT). Over the depth-integrated euphotic zone, mesozooplankton accounted for 30% of the combined grazing losses of phytoplankton to micro- and meso-herbivores in Opal, as compared to 13% at control stations. Estimates of active export flux by migrating zooplankton averaged 0.81 mmol C m⁻² d⁻¹ in Cyclone Opal and 0.37 mmol C m⁻² d⁻¹ at OUT stations, 53% and 24%, respectively, of the carbon export measured by passive sediment traps. Migrants also exported 0.18 mmol N m⁻² d⁻¹ (117% of trap N flux) in Cyclone Opal compared to 0.08 mmol N m⁻² d⁻¹ (51% of trap flux) at control stations. Overall, the food-web importance of mesozooplankton increased in Cyclone Opal both in absolute and relative terms. Diel migrants provided evidence for enhanced export flux in the eddy that was missed by sediment trap and ²³⁴Th techniques, and migrant-mediated flux was the major export term in the observed bloom-perturbation response and N mass balance of the eddy.     

The recovery of microalgal production and biomass in a South African temporarily open/closed estuary, following mouth breaching

Mouth breaching is a recurrent event in temporarily open/closed estuaries (TOCEs). Such disturbances result in flushing and sediment scouring, reducing the microalgal biomass stock. The depletion of these microalgae may have negative repercussions in the form of depleted stocks of commercial fish, game fish, crustaceans and mollusks. The aim of this investigation was therefore: (1) to monitor the recovery of microalgal biomass and production following a breaching event; and (2) to determine the key environmental parameters influencing primary production during the open and recovery phases. Phytoplankton and benthic microalgal production was measured (¹⁴C-uptake method) successively during the closed, open and recovery phases of the Mdloti TOCE (South Africa). Upon breaching, 94–99% of microalgal biomass was washed out to sea through flushing and sediment scouring. A temporary recovery of phytoplankton and benthic microalgal biomass was observed during the open phase, but this was not sustained because of continual flushing and scouring of the sediment. During the re-closure (recovery phase), microalgal biomass immediately increased, reaching pre-breaching levels 35–40 days following the breaching event. In contrast to biomass, autochthonous pelagic primary production reached a maximum level (341 mg C m⁻² h⁻¹) during the open phase. Pelagic primary production normalized to biomass (P^B) significantly increased during the open phase. This is attributed to a favorable combination of optimum light conditions, high influx of macronutrients and high water temperatures (33 °C). Similarly, benthic primary production normalized to biomass (P^B) peaked during the open phase (35 mg C mg chl-a⁻¹ h⁻¹). Multivariate analysis showed that major variations in primary production were mainly controlled by temperature, dissolved inorganic nitrogen (DIN) to phosphorus (DIP) molar ratios (water-column and pore-water) and light extinction (K_d), all of which were regulated by the state of the mouth.     

A depocenter of organic matter at 7800 m depth in the SE Pacific Ocean

The Atacama trench, the deepest ecosystem of the southern Pacific Ocean (ca. 8000 m depth) was investigated during the Atacama Trench International Expedition. Sediments, collected at three bathyal stations (1040–1355 m depth) and at a hadal site (7800 m) were analyzed for organic matter quantity and biochemical composition (in terms of phytopigments, proteins, carbohydrates and lipids), bacterial abundance, biomass and carbon production and extracellular enzymatic activities. Functional chlorophyll-a (18.0±0.10 mg m⁻²), phytodetritus (322.2 mg m⁻²) and labile organic carbon (16.9±4.3 g C m⁻²) deposited on surface sediments at hadal depth (7800 m) reached concentrations similar to those encountered in highly productive shallow coastal areas. High values of bacterial C production and aminopeptidase activity were also measured (at in situ temperature and 1 atm). The chemical analyses of the Atacama hadal sediments indicate that this trench behaves as a deep oceanic trap for organic material. We hypothesize that, despite the extreme physical conditions, benthic microbial processes might be accelerated as a result of the organic enrichment.     

Impact of freshwater influx on microzooplankton mediated food web in a tropical estuary (Cochin backwaters – India)

The diversity, abundance and biomass of microzooplankton in Cochin backwaters were studied for the first time during pre-summer monsoon to peak of summer monsoon (April–July 2003) to understand the impact of large freshwater influx. Microzooplankton abundance and biomass were highest during pre-summer monsoon (av. 3817 ind. L⁻¹ and 146 μg C L⁻¹) that declined with the onset (av. 2052 ind. L⁻¹ and 45 μg C L⁻¹) and peak (av. 409 ind. L⁻¹ and 10 μg C L⁻¹) summer monsoon. Species diversity, richness and evenness of microzooplankton also showed similar trends as that of abundance and biomass. Grazing experiment showed that microzooplankton consumes 43 ± 1% of the daily phytoplankton standing stock during the high saline condition (27.5). Low abundance of microzooplankton during summer monsoon period (1/8 of the pre-summer monsoon value) along with the concomitant occurrence of low mesozooplankton (1/8 times of pre-summer monsoon value) suggests that there could be a general lack of planktonic grazers. This would result in a weak transfer of primary and bacterial carbon to higher trophic levels, eventually leaving behind much unconsumed basic food in the estuary during summer monsoon. Thus a major portion of the primary carbon either settles down or gets transported to the coastal regions during monsoon. High flushing of Cochin backwaters also facilitates faster removal of primary producers to the coastal regions during monsoon.     

Photosynthate allocation in a temperate sea over an annual cycle: the relationship between protein synthesis and phytoplankton physiological state

The seasonal and vertical variations in the patterns of photosynthate allocation into biomolecules by natural phytoplankton assemblages were determined, together with their species composition, in a coastal station of the central Cantabrian Sea (southern Bay of Biscay). Chlorophyll-a concentration ranged from values below 20 mg m⁻² in winter to values above 80 mg m⁻² during spring and during an upwelling event in summer. Low primary production rates (<300 mgC m⁻² d^-1) were measured during winter and during summer stratification periods. The rate of C fixation during summer upwelling conditions exceeded 3500 mgC m⁻² d⁻¹. In terms of photosynthate partitioning, proteins were the dominant fraction, as they typically accounted for >30% of total photo-assimilated C, with polysaccharides and low molecular weight metabolites showing incorporation percentages around 10–30%. Relative C incorporation into lipids was generally <15%. Recurrent patterns of vertical variability in photosynthate partitioning were observed: the relative synthesis of proteins increased toward the bottom of the euphotic zone, whereas the relative C incorporation into polysaccharides and lipids tended to be higher near the surface. When primary production decreased, the synthesis of proteins was maintained more than that of other molecules. Throughout the year, the relative synthesis of proteins was inversely correlated with phytoplankton biomass, production and growth rate. The conservation of protein synthesis under growth-limiting conditions and the enhancement of lipid and polysaccharide synthesis when irradiance is high seem to constitute general patterns of photosynthate partitioning in marine phytoplankton. In our study, these patterns represented metabolic strategies of phytoplankton in response to changing environmental factors, rather than the effect of variations in the species composition of the community.     

Abundance, variability, and potential grazing impact of planktonic ciliates in the open subaratic Pacific Ocean

The abundance and variability of planktonic ciliates in the open subarctic Pacific were determined during four month-long cruises in 1987 and 1988. The ciliate community, numerically dominated by relatively small aloricate choreotrichs, was comparable in abundance to communities in a range of oceanic and neritic environments, including waters with much higher average chlorophyll concentrations. Integrated (0–80m) ciliate biomass was typically 100–200mgC m⁻², although 3- to 4-fold higher levels were observed on two occasions in spring. Ciliate community biomass, in general, was dominated by large (>20μm width) individuals, although in August 1988 the biomass of smaller cells was as great or greater. The estimated grazing impact of the ciliate community averaged 20% of the primary production. On one instance in May 1988, however, a large biomass of ciliates led to an estimated grazing impact equivalent to 55% of phytoplankton production. While ciliates may be major phytoplankton grazers during sporadic ciliate “blooms”, dino- and other heterotrophic flagellates, which make up the bulk of microheterotroph biomass, must normally be of equal or greater importance as herbivores in this ocean region.     

Does transported seagrass provide an important trophic link in unvegetated, nearshore areas?

The contribution of detritus from seagrass and other primary producers to faunal production in unvegetated nearshore areas was examined primarily using stable isotopes. Fish, macroinvertebrates, meiofauna and primary producers (seagrasses, macroalgae, seston and benthic microalgae) were sampled from sites in south-western Australia. All samples were analysed for δ¹³C and δ¹⁵N values and fish gut contents were determined. δ¹³C values for seagrasses in the region were high compared to other macrophytes, ranging from 49.9 to −8.2‰ compared to −19.8 to −12.6‰ for macroalgae. The δ¹⁵N values ranged between 4.0 and 7.7‰ for the red, brown and green algae, and between 3.2 and 5.9‰ for seagrasses. Seston and benthic microalgae samples had a mean δ¹³C value of −12.8 and −14.0‰, respectively, and their δ¹⁵N values were comparable to the macroalgae. All invertebrate fauna had mean δ¹³C values considerably lower than seagrasses. However, individual samples harpacticoid copepods and polychaetes had a value as high as −11.7‰. δ¹⁵N values for consumers were higher than those of the primary producers, except for copepods and amphipods. The δ¹³C values for fish had a relatively small range, between −16.6 and −13.1‰, and the δ¹⁵N values of fish were elevated compared to the invertebrates and primary producers, ranging mostly between 10.0 and 12.6‰. Mixing model analysis based on δ¹³C values indicated that seagrass ranked low as a likely carbon source for all invertebrates other than harpacticoid copepods at a single site and some samples of polychaetes. The δ¹³C values for fish were similar to those of a combination of harpacticoid and calanoid copepods, amphipods and polychaetes. The consumption of harpacticoid copepods by some fish species indicates that Amphibolis and Posidonia species in south-western Australia can contribute to the food web of unvegetated nearshore areas as detritus, but brown algae is likely to make a greater contribution. At least for the time of year that was sampled, the flow of detrital seagrass material into the foodweb may be mediated by specific detrivores, in this case harpactacoid copepods, rather than by all detritivores.     

Factors influencing benthic bacterial abundance, biomass, and activity on the northern continental margin and deep basin of the Gulf of Mexico

As part of a larger project on the deep benthos of the Gulf of Mexico, an extensive data set on benthic bacterial abundance (n>750), supplemented with cell-size and rate measurements, was acquired from 51 sites across a depth range of 212–3732 m on the northern continental slope and deep basin during the years 2000, 2001, and 2002. Bacterial abundance, determined by epifluorescence microscopy, was examined region-wide as a function of spatial and temporal variables, while subsets of the data were examined for sediment-based chemical or mineralogical correlates according to the availability of collaborative data sets. In the latter case, depth of oxygen penetration helped to explain bacterial depth profiles into the sediment, but only porewater DOC correlated significantly (inversely) with bacterial abundance (p<0.05, n=24). Other (positive) correlations were detected with TOC, C/N ratios, and % sand when the analysis was restricted to data from the easternmost stations (p<0.05, n=9–12). Region-wide, neither surface bacterial abundance (3.30–16.8×10⁸ bacteria cm⁻³ in 0–1 cm and 4–5 cm strata) nor depth-integrated abundance (4.84–17.5×10¹³ bacteria m⁻², 0–15 cm) could be explained by water depth, station location, sampling year, or vertical POC flux. In contrast, depth-integrated bacterial biomass, derived from measured cell sizes of 0.027–0.072 μm³, declined significantly with station depth (p<0.001, n=56). Steeper declines in biomass were observed for the cross-slope transects (when unusual topographic sites and abyssal stations were excluded). The importance of resource changes with depth was supported by the positive relationship observed between bacterial biomass and vertical POC flux, derived from measures of overlying productivity, a relationship that remained significant when depth was held constant (partial correlation analysis, p<0.05, df=50). Whole-sediment incubation experiments under simulated in situ conditions, using ³H-thymidine or ¹⁴C-amino acids, yielded low production rates (5–75 μg C m⁻² d⁻¹) and higher respiration rates (76–242 μg C m⁻² d⁻¹), with kinetics suggestive of resource limitation at abyssal depths. Compared to similarly examined deep regions of the open ocean, the semi-enclosed Gulf of Mexico (like the Arabian Sea) harbors in its abyssal sediments a greater biomass of bacteria per unit of vertically delivered POC, likely reflecting the greater input of laterally advected, often unreactive, material from its margins.     

Trophic Status of the South-Eastern Baltic Sea: A Comparison of Coastal and Open Areas

Primary production, nutrient concentrations, phytoplankton biomass (incl. chlorophyll a) and water transparency (Secchi depth), are important indicators of eutrophication. Earlier basin-wide primary production estimates for the Baltic Sea, a shallow shelf sea, were based mainly on open-sea data, neglecting the fundamentally different conditions in the large river plumes, which might have substantially higher production. Mean values of the period 1993–1997 of nutrient concentrations (phosphate, nitrate, ammonium and silicate), phytoplankton biomass, chlorophyll a (chl a) concentration, turbidity and primary production were calculated in the plumes of the rivers Oder, Vistula and Daugava and Klaipeda Strait as well as the open waters of the Arkona Sea, Bornholm Sea, eastern Gotland Sea and the Gulf of Riga. In the plumes, these values, except for primary production, were significantly higher than in the open waters. N:P ratios in the plumes were >16 (with some exceptions in summer and autumn), indicating potential P-limitation of phytoplankton growth, whereas they were <16 in the open Baltic Proper, indicating potential N-limitation. On the basis of in situ phytoplankton primary production, phytoplankton biomass and nutrient concentrations, the large river plumes and the Gulf of Riga could be characterized as eutrophic and the outer parts of the coastal waters and the open sea as mesotrophic. Using salinity to define the border of the plumes, their mean extension was calculated by means of a circulation model. Taking into account the contribution of coastal waters, the primary production in the Baltic Proper and the Gulf of Riga was 42·6 and 4·3×10⁶ t C yr⁻¹, respectively. Hence, an annual phytoplankton primary production in the whole Baltic Sea was estimated at 62×10⁶ t C yr⁻¹. The separate consideration of the plumes had only a minor effect on the estimation of total primary production in comparison with an estimate based on open sea data only. There is evidence for a doubling of primary production in the last two decades. Moreover, a replacement of diatoms by dinoflagellates during the spring bloom was noticed in the open sea but not in the coastal waters. A scheme for trophic classification of the Baltic Sea, based on phytoplankton primary production and biomass, chl a and nutrient concentrations, is proposed.     

Modeling environmental effects on the size-structured energy flow through marine ecosystems. Part 2: Simulations

Numerical simulations using a physiologically-based model of marine ecosystem size spectrum are conducted to study the influence of primary production and temperature on energy flux through marine ecosystems. In stable environmental conditions, the model converges toward a stationary linear log–log size-spectrum. In very productive ecosystems, the model predicts that small size classes are depleted by predation, leading to a curved size-spectrum.It is shown that the absolute level of primary production does not affect the slope of the stationary size-spectrum but has a nonlinear effect on its intercept and hence on the total biomass of consumer organisms (the carrying capacity). Three domains are distinguished: at low primary production, total biomass is independent from production changes because loss processes dominate dissipative processes (biological work); at high production, ecosystem biomass is proportional to primary production because dissipation dominates losses; an intermediate transition domain characterizes mid-production ecosystems. Our results enlighten the paradox of the very high ecosystem biomass/primary production ratios which are observed in poor oceanic regions. Thus, maximal dissipation (least action and low ecosystem biomass/primary production ratios) is reached at high primary production levels when the ecosystem is efficient in transferring energy from small sizes to large sizes. Conversely, least dissipation (most action and high ecosystem biomass/primary production ratios) characterizes the simulated ecosystem at low primary production levels when it is not efficient in dissipating energy.Increasing temperature causes enhanced predation mortality and decreases the intercept of the stationary size spectrum, i.e., the total ecosystem biomass. Total biomass varies as the inverse of the Arrhenius coefficient in the loss domain. This approximation is no longer true in the dissipation domain where nonlinear dissipation processes dominate over linear loss processes. Our results suggest that in a global warming context, at constant primary production, a 2–4 °C warming would lead to a 20–43% decrease of ecosystem biomass in oligotrophic regions and to a 15–32% decrease of biomass in eutrophic regions.Oscillations of primary production or temperature induce waves which propagate along the size-spectrum and which amplify until a “resonant range” which depends on the period of the environmental oscillations. Small organisms oscillate in phase with producers and are bottom-up controlled by primary production oscillations. In the “resonant range”, prey and predators oscillate out of phase with alternating periods of top-down and bottom-up controls. Large organisms are not influenced by bottom-up effects of high frequency phytoplankton variability or by oscillations of temperature.     

Major contribution from mesopelagic plankton to heterotrophic metabolism in the upper ocean

Measurements of plankton respiration and heterotrophic bacterial abundance and production were made at seven deep water stations within the upper 500 m of the Gulf of Mexico during the summer of 1995. Bacterial abundance [(1.1–4.6)×108 1⁻¹] and rates of bacterial production (2–19 nM C h⁻¹) and plankton respiration (50–245 nM O₂ h⁻¹) decreased with depth by four- to nine-fold, and were similar to those reported for oligotrophic waters. Bacterial turnover times increased with depth from approximately 1 to 5 days. Bacterial growth efficiencies decreased from 15% at the surface to 8% at 500 m. Depth-integrated plankton respiration exceeded known estimates of primary production for the region, suggesting that heterotrophic utilization of previously and concurrently produced organic matter (e.g. spring phytoplankton growth, and summer blooms of Trichodesmium sp.) was occurring during the summer. Estimates for the upper 500 m showed that roughly half of the bacterial biomass (56%), bacterial production (49%), and plankton respiration (60%) occurred below the euphotic zone. Routine oceanographic studies have focused exclusively on the metabolic activity occurring within the euphotic zone. Our measurements, however, indicate that mesopelagic plankton also contribute substantially to heterotrophic metabolism and nutrient cycling in the ocean.     

Size and taxonomic plankton community structure and carbon flow at the equator, 175‡E during 1990–1994

Size and taxonomic structure of plankton community carbon biomass for the 0.2–2000 μm equivalent spherical diameter range were determined at the equator at 175°E in September 1990–1993 and April 1994. Total biomass of the plankton community ranged from 1944 to 3448 mg C m⁻². Phytoplankton, zooplankton and bacteria carbon biomasses were 604–1669 mg C m^-2, 300–797 mg C m², and 968–1200 mg C m^-2, and the percentages were 31–54%, 15–26%, and 29–54%, respectively. Biomass of heterotrophic bacteria was always the largest fraction andProchlorococcus biomass was second. Heterotrophic and autotrophic flagellates and dinoflagellates in the nanoplankton size range and copepods (adults and copepodites) in the mesoplankton range were also high. Relatively small biomass was observed in the microplankton size range. The differences in integrated biomass of plankton community for El Nin˜o type oligotrophic conditions of September 1990–1993 and non-El Nifio type mesotrophic conditions of April 1994 were generally small compared with the interannual difference during 1990–1993. However, the percentage ofProchlorococcus in phytoplankton carbon biomass was larger in non-El Nin˜o year. Biomasses of cyanobacteria, diatom, dinoflagellates, nauplii of copepods, and crustaceans other than copepods were larger in the non-El Nin˜o year. Primary production increased significantly from El Nin˜o to non-El Nin˜o years. Carbon flow through the plankton food chain was estimated using the plankton carbon biomass data, primary production measurements, and published empirical relationships.     

Amphipod prey of gray whales in the northern Bering Sea: Comparison of biomass and distribution between the 1980s and 2002–2003

The ampeliscid amphipod community in the Chirikov Basin of the northern Bering Sea was a focus of study during the 1980s because they were a major food for the Eastern North Pacific (ENP) population of gray whales Eschrichtius robustus. Information from the 1980s benthic investigations, published accounts of ENP gray whale population trends and the occurrence in 1999–2000 of an unusual number of gray whale mortalities prompted concern that the whale population may have exceeded the carrying capacity of its food base. Therefore, during two cruises per year between June and September, 2002 and 2003, we resampled the 20 stations occupied during the 1980s, to determine if there had been any significant changes in ampeliscid abundance and biomass. During 2002–2003, average ampeliscid dry weight biomass was about 28±10 g m⁻² (95% confidence interval), a decline of nearly 50% from maximum values in the 1980s. Amphipod length measurements indicated that the declines were due mainly to the absence of the larger animals (20–30 mm length). Two hypotheses were considered regarding the amphipod declines: gray whale predation and climate. Ampeliscid production (105 kcal m⁻² yr⁻¹) and gray whale energy requirements (1.6×10⁸ kcal individual⁻¹ yr⁻¹) indicated that as little as 3–6% of the current estimate of the ENP gray whale population could remove 10–20% of the annual ampeliscid production from the study site in 2002–2003, a finding consistent with the hypothesis that top-down control by foraging whales was the primary cause of the observed declines. A 10-yr time series of temperature near the bottom in the Bering Strait and northward transport did not reveal a consistent trend between 1990 and 2001, suggesting that climate influences were not the major cause of the observed declines. Arctic ampeliscids have slow growth rates and long generation times; therefore the ampeliscid community may require decades to recover to densities observed in the 1980s. Predicted warming trends in the northern Bering Sea could impact ampeliscid recovery by lowering primary production or altering the community composition of the benthos.     

Secondary production of Ampelisca mississippiana Soliman and Wicksten 2007 (Amphipoda, Crustacea) in the head of the Mississippi Canyon, northern Gulf of Mexico

Annual production was calculated for the dominant ampeliscid amphipod Ampelisca mississippiana [Soliman, Y., Wicksten, M., 2007. Ampelisca mississippiana a new species (Amphipoda: Gammaredea) dominated the head of the Mississippi Canyon (Northern Gulf of Mexico). Zootaxa, submitted] at the head of the Mississippi Canyon in the northern Gulf of Mexico. Average densities were 12,094±2499 ind m⁻², with secondary production of 6.93 g dry wt m⁻² yr⁻¹, based on the “size-frequency method” [Hynes-Hamilton, H.B.N., Coleman, M., 1968. A simple method for assessing the annual production of stream benthos. Limnology and Oceanography 13, 569–573; Menzies, C.A., 1980. A note on the Hynes-Hamilton method of estimating secondary production. Limnology and Oceanography 25(4), 770–773], with a production/biomass (P/B) ratio of 3.11. Growth rates of this magnitude are comparable to available data for freshwater and shallow marine ampeliscids, but are unexpectedly high for deep-ocean habitats. Growth efficiency appeared to be approximately 35% (Growth/Assimilation×100).