Vertical distribution of zooplankton and active flux across an anticyclonic eddy in the Canary Island waters

The vertical distribution (0–900 m) of zooplankton biomass and indices of feeding (gut fluorescence, GF) and metabolism (electron transfer system, ETS) were studied across an anticyclonic eddy south of Gran Canaria Island (Canary Islands). Two dense layers of organisms were clearly observed during the day, one above 200 m and the other at about 500 m, coincident with the deep scattering layer (DSL). The biomass displacement due to interzonal migrants in the euphotic zone was more than 2-fold higher than that previously reported for the southern area of this archipelago. The gut flux estimated (0.14–0.44 mgC m⁻² d⁻¹) was similar to the values previously found in the Canaries. The respiratory flux outside the eddy (1.85 mgC m⁻² d⁻¹) was in the lower range of values reported for this area. Inside the eddy, migrant biomass and respiration rates were 2- and 4- fold higher than in the surrounding waters. Active flux mediated by diel vertical migrants inside the eddy (8.28 mgC m⁻² d⁻¹) was up to 53% of the passive carbon flux to the mesopelagic zone (15.8 mgC m⁻² d⁻¹). It is, therefore, suggested that the anticyclonic eddy enhanced both migration from deep waters and active flux.     

The supply of nutrients due to vertical turbulent mixing: A study at the Porcupine Abyssal Plain study site in the northeast Atlantic

As part of a multidisciplinary cruise to the Porcupine Abyssal Plain (PAP) study site (49°00′N 16°30′W), in June and July 2006, observations were made of the vertical nitrate flux due to turbulent mixing. Daily profiles of nitrate and turbulent mixing, at the central PAP site, give a mean nitrate flux into the euphotic zone of 0.09 (95% confidence intervals: 0.05–0.16) mmol N m⁻² d⁻¹. This is a factor of 50 lower than the mean observed rate of nitrate uptake within the euphotic zone (5.1±1.3 mmol N m⁻² d⁻¹). By using our direct observations to ‘validate’ a previously published parameterisation for turbulent mixing, we further quantify the variability in the vertical turbulent flux across a roughly 100×100 km region centred on the PAP site, using hydrographic data. The flux is uniformly low (0.08±0.26 mmol N m⁻² d⁻¹, the large standard deviation being due to a strongly non-Gaussian distribution) and is consistent with direct measurements at the central site. It is demonstrated that on an annual basis convective mixing supplies at least 40-fold more nitrate to the euphotic zone than turbulent mixing at this location. Other processes, such as those related with mesoscale phenomena, may also contribute significantly.     

High primary productivity and f-ratio in summer in the Ulleung basin of the East/Japan Sea

To better understand the cause of high summer primary productivity in the Ulleung Basin located in the southwest part of the East/Japan Sea, the spatial dynamics of primary, new, and regenerated productivities (PP, NP, and RP) were examined along the path of the Tsushima Warm Current system in summer 2008. We compared hydrographic and chemical parameters in the Ulleung Basin with those of the Kuroshio Current in the Western Pacific Ocean and the East China Sea. In summer, integrated primary productivity (IPP, 0.37–0.96 g C m⁻² d⁻¹) and integrated new productivity (INP, 26–221 mg N m⁻² d⁻¹) within the euphotic zone in the Ulleung Basin were higher than those in the East China Sea and the Western Pacific Ocean (0.17–0.28 g C m⁻² d⁻¹, 2−5 mg N m⁻² d⁻¹, respectively). In contrast, there was no pronounced spatial variation in integrated regenerated productivity (IRP, 43–824 mg N m⁻² d⁻¹). Strong positive correlations between IPP and INP (also the f-ratio), and between nitrate uptake rate in the mixed layer and nitrate upward flux through the top of pycnocline in summer in the Ulleung Basin imply that the high IPP was mainly supported by supply of nitrate from the underlying water in the euphotic zone. Shallowing of the pycnocline depth as the current enters the East/Japan Sea facilitates nitrate supply from the nutrient-replete cold water immediately below the pycnocline through nitrate upward flux. A subsurface maximum in PP at or above the pycnocline and a high f-ratio further support the importance of this source of nitrate for maintaining the high summer PP in the Ulleung Basin. In comparison, the high PP layer was observed at the surface in the following fall and spring in the Ulleung Basin. Our results demonstrate the importance of hydrographic features in enhancing PP in this oligotrophic Tsushima Warm Current system.     

Consumption and production on scales of a few days of inorganic carbon,nitrate and oxygen by the planktonic community: results of continuous measurements at the Dyfamed Station in the northwestern Mediterranean Sea (May 1995)

Continuous measurements between 0 and 200 m depth were performed every 2 h over two separate periods of four days at a station in the open northwestern Mediterranean Sea (Dyfamed Station) during the Dynaproc cruise in May 1995. Estimates of the daily variations in profiles of temperature, partial pressure of CO₂, oxygen, chlorophyll a and nutrients were obtained. The distributions of the various physical and chemical properties were clearly different during the two time series, which were separated by a period of 11 days during which a wind event occurred. The mean daily utilization or production due to biological processes of dissolved inorganic carbon (DIC), nitrate+nitrite and oxygen were calculated along isopycnals using a vertical diffusion model. Between the surface and about 20 m depth, DIC was consumed and O₂ released during the two time series while the nitrate+nitrite concentrations as well as supplies were zero. After the wind event, the O₂ : C : N ratios of consumption (or production) were, on average, near the Redfield ratios, but during the first time series, the C : N utilization ratio between 20 and 35 m was two to three times that of Redfield stoichiometry and the oxygen release was low. The integrated net community production (NCP) in terms of carbon was equivalent during the two time series, whereas the chlorophyll a biomass was twice as high, on average, during the first time series but did decrease. These results imply that the production systems were different during the two periods. The first time series corresponds to a period at the end of production, due to the nutrient depletion in the euphotic layer. The formation of degradation products of the living material in dissolved organic form is probably important as indicated by the high C : N utilization ratios. The second time series corresponds to a reactivation of the primary production due to the upward shift of nutrients after the wind event.     

Community structure and photosynthetic physiology of phytoplankton in the northwest subarctic Pacific during an in situ iron fertilization experiment (SEEDS-II)

Temporal changes in the abundance, community composition, and photosynthetic physiology of phytoplankton in surface waters were investigated during the second in situ iron (Fe) fertilization experiment in the NW subarctic Pacific (SEEDS-II). Surface chlorophyll a concentration was 0.75 mg m⁻³ on the day before the first Fe enrichment (i.e. Day 0), increased ca. 3-fold until Day 13 after two Fe additions, and thereafter declined with time. The photochemical quantum efficiency (F_v/F_m) and functional absorption cross-section (σPSII) of photosystem II for total phytoplankton in surface waters increased and decreased inside the Fe-enriched patch through Day 13, respectively. These results indicate that the photosynthetic physiological condition of the phytoplankton improved after the Fe infusions. However, the maximum F_v/F_m value of 0.43 and the maximum quantum yield of carbon fixation (φ_max) of 0.041 mol C (mol photon)⁻¹ during the development phase of the bloom were rather low, compared to their theoretical maximum of ca. 0.65 and 0.10 mol C (mol photon)⁻¹, respectively. Diatoms, which were mainly composed of oceanic species, did not bloom, and autotrophic nanoflagellates such as cryptophytes and prasinophytes became predominant in the phytoplankton community inside the Fe-enriched patch. In ferredoxin/flavodoxin assays for micro-sized (20–200 μm in cell length) diatoms, ferredoxin was not detected but flavodoxin expressions consistently occurred with similar levels both inside and outside the Fe-enriched patch, indicating that the large-sized diatoms were stressed by Fe bioavailability inside the Fe-enriched patch even after the Fe enrichments. Our data suggest that the absence of a Fe-induced large-sized diatom bloom could be partly due to their Fe stress throughout SEEDS-II.     

Thermocline ventilation and oxygen utilization rates in the subtropical North Pacific based on CFC distributions during WOCE

Thermocline ventilation rates for the subtropical North Pacific are determined using a 1-dimensional (meridional) along-isopycnal advective–diffusive model tuned to chlorofluorocarbon (CFC) concentrations measured along 152°W in 1991 during WOCE P16. Mean southward advection rates in the subtropics range from 1.03 to 0.56 cm s^-1 between σ_θ=25.5 and 26.6. Model-derived ventilation times for the subtropical gyre increase from about 10 to 27 years for that isopycnal range. Oxygen utilization rates (OURs) determined using the advective-diffusive model decrease with depth from 6.6 to 3.2 μmol kg^-1 yr^-1 between σ_θ=25.5 and 26.6. Extrapolation of the OUR versus depth trend to the base of the euphotic zone with the 1/Z power function of Martin et al. (1987) and integration from 500 to 100 m depth implies a carbon export rate from the overlying euphotic zone of 2.2±0.5 moles C m^-2 yr^-1 at 30°N, 152°W. Analysis of the WOCE radiocarbon and salinity distributions indicates that zonal and cross-isopycnal transport terms would have to be considered in modeling these tracers in the subtropical North Pacific.     

Latitudinal distribution of microbial plankton abundance,production, and respiration in the Equatorial Atlantic in autumn 2000

Phytoplankton and bacterial abundance, size-fractionated phytoplankton chlorophyll-a (Chl-a) and production together with bacterial production, microbial oxygen production and respiration rates were measured along a transect that crossed the Equatorial Atlantic Ocean (10°N–10°S) in September 2000, as part of the Atlantic Meridional Transect 11 (AMT 11) cruise. From 2°N to 5°S, the equatorial divergence resulted in a shallowing of the pycnocline and the presence of relatively high nitrate (>1 μM) concentrations in surface waters. In contrast, a typical tropical structure (TTS) was found near the ends of the transect. Photic zone integrated ¹⁴C primary production ranged from ∼200 mg C m⁻² d⁻¹ in the TTS region to ∼1300 mg C m⁻² d⁻¹ in the equatorial divergence area. In spite of the relatively high primary production rates measured in the equatorial upwelling region, only a moderate rise in phytoplankton biomass was observed as compared to nearby nutrient-depleted areas (22 vs. 18 mg Chl-a m⁻², respectively). Picophytoplankton were the main contributors (>60%) to both Chl-a biomass and primary production throughout the region. The equatorial upwelling did not alter the phytoplankton size structure typically found in the tropical open ocean, which suggests a strong top-down control of primary producers by zooplankton. However, the impact of nutrient supply on net microbial community metabolism, integrated over the euphotic layer, was evidenced by an average net microbial community production within the equatorial divergence (1130 mg C m⁻² d⁻¹) three-fold larger than net production measured in the TTS region (370 mg C m⁻² d⁻¹). The entire region under study showed net autotrophic community metabolism, since respiration accounted on average for 51% of gross primary production integrated over the euphotic layer.     

Microbial respiration in the Levantine Sea: evolution of the oxidative processes in relation to the main Mediterranean water masses

First data on microbial respiration in the Levantine Sea are reported with the aim of assessing the distribution of oxidative processes in association with the main Mediterranean water masses and the changing physical structure determined by the Eastern Mediterranean Transient. Respiratory rates, in terms of metabolic carbon dioxide production, were estimated from measured electron transport system activities in the polygonal area of the Levantine Sea (32.5–36.5 N Latitude, 26.0–30.25 E Longitude) and at Station Geo’95, in the Ionian Sea (35°34.88 N; 17°14.99 E). At the Levantine Sea, the mean carbon dioxide production rate decreased from the upper to the deeper layers and varied from 22.0±12.4 μg C h⁻¹ m⁻³ in the euphotic layer to 1.30±0.5 μg C h⁻¹ m⁻³ in the depth range between 1600 and 3000 m. Significant differences were found among upper, intermediate and bottom layers. The euphotic zone supported a daily carbon dioxide production of 96.6 mg C d⁻¹ m⁻² while the aphotic zone (between 200 and 3000 m) sustained a 177.1 mg C d⁻¹ m⁻² carbon dioxide production. In Station Geo’95, the carbon dioxide production rates amounted to 170.4 and 102.2 mg C d⁻¹ m⁻² in the euphotic and aphotic zones, respectively. The rates determined in the identified water masses showed a tight coupling of respiratory processes and Mediterranean circulation patterns. The increasing respiratory rates in the deep layers of the Levantine Sea are explained by the introduction of younger waters recently formed in the Aegean Sea.     

New and regenerated production in the Almeria-Oran front area,eastern Alboran Sea

Uptake and regeneration of nitrogen in the Almeria-Oran frontal zone (SW Mediterranean) and adjacent (Atlantic and Mediterranean) systems were studied during the Almofront I cruise (JGOFS-France). The frontal zone was characterized by an upsloping of nitracline from about 50 m in the adjacent systems to 25–30 m within. Along with nitrate, ammonium, chlorophyll a and particulate organic nitrogen also were at higher concentrations in the frontal zone than in the adjacent waters.The nitrate uptake rates were significantly higher in the frontal zone (up to 6.4 nmol l⁻¹ h⁻¹) than in the Atlantic and Mediterranean waters (generally <1 nmol l⁻¹ h⁻¹) indicating a significant increase of new production at the front. This increase was related to the upsloping of the nitracline as shown by the significant correlation (p<0.05) between new production and depth of the nitracline. The new production in the Almeria-Oran was much lower than those recorded in other oceanic and coastal fronts. This could be related to the fact that the nitracline did not rise up to the surface and the high concentrations of nitrate were confined to deeper layers where the ambient light intensity was less. Nitrate uptake in the frontal zone was significantly higher, by 1.7–5.8 times (average 4.2), than the calculated diffusive flux of nitrate, suggesting that vertical advection may be an important source of nitrate. New production rates at the front were also significantly higher (3–9 times, average 5.8) than the PON flux to 100 m depth estimated by sediments traps (Journal of Marine Systems 5, 377–389), suggesting a strong decoupling between surface production and downward flux of POM in the frontal zone.The north–south gradient observed with different parameters indicates the presence of a transfrontal secondary circulation. This distribution also suggests that the primary production in the front is initially nitrate-based, with a diatom-herbivore food chain, whereas regenerated production, associated with an intense recycling of organic matter, later becomes progressively important in time and space.     

Diapycnal nutrient fluxes on the northern boundary of Cape Ghir upwelling region

In this study we estimate diffusive nutrient fluxes in the northern region of Cape Ghir upwelling system (Northwest Africa) during autumn 2010. The contribution of two co-existing vertical mixing processes (turbulence and salt fingers) is estimated through micro- and fine-structure scale observations. The boundary between coastal upwelling and open ocean waters becomes apparent when nitrate is used as a tracer. Below the mixed layer (56.15±15.56 m), the water column is favorable to the occurrence of a salt finger regime. Vertical eddy diffusivity for salt (K_s) at the reference layer (57.86±8.51 m, CI 95%) was 3×10⁻⁵ (±1.89×10⁻⁹, CI 95%) m² s⁻¹. Average diapycnal fluxes indicate that there was a deficit in phosphate supply to the surface layer (6.61×10⁻⁴ mmol m⁻² d⁻¹), while these fluxes were 0.09 and 0.03 mmol m⁻² d⁻¹ for nitrate and silicate, respectively. There is a need to conduct more studies to obtain accurate estimations of vertical eddy diffusivity and nutrient supply in complex transitional zones, like Cape Ghir. This will provide us with information about salt and nutrients exchange in onshore–offshore zones.     

A method for the determination of nitrification rates in oligotrophic marine seawater by gas chromatography/mass spectrometry

An isotope dilution method has been developed to determine by gas chromatography/mass spectrometry (GC/MS) the rates of ammonium and nitrite oxidation in severely oligotrophic marine waters. The method is based on the formation of sudan-1 from nitrite, or from nitrate following reduction to nitrite. Samples were collected by solid phase extraction and purified by high performance liquid chromatography (HPLC). A deuterated sudan-1 internal standard was synthesized, purified by HPLC and used for quantitative analysis. Concentrations of NO₂⁻ and NO₃⁻ were generally < 2 nmol/kg and < 5 nmol/kg respectively, typical of oligotrophic surface waters, and turnover times for the inorganic N pools ranged from < 1 day to > 10 days. Significant rates of nitrification were measured in the surface oligotrophic ocean, with rates of ammonium and nitrite oxidation generally within the range of 10–500 pmol/kg/h. Consequently, a significant proportion of daily NO₃⁻ assimilation by marine phytoplankton is regenerated, and not new. In a case study of the oligotrophic gyre of the North Atlantic, the influence of NH₄⁺ regeneration and nitrification on f-ratio values suggests that in the oligotrophic ocean, f-ratio values may be significantly, and sometimes grossly, overestimated.     

Physical-biological coupling in the Algerian Basin (SW Mediterranean): Influence of mesoscale instabilities on the biomass and production of phytoplankton and bacterioplankton

The biomass and production of phytoplankton and bacterioplankton was investigated in relation to the mesoscale structures found in the Algerian Current during the ALGERS'96 cruise (October 1996). Biological determinations were carried out in three transects between 0° and 2°E aimed at crossing a so-called event, formed by a coastal anticyclonic eddy associated with an offshore cyclonic eddy to the west. The concentration of chlorophyll a (Chl) was maximum (>1.2 mg m⁻³) within the cyclonic eddy and at the frontal zones between the Modified Atlantic Water (MAW) of the Algerian Current and the Mediterranean waters further north. Chl (total and >2 μm) was significantly correlated with proxies of nutrient flux into the upper layers. Autotrophic picoplankton and heterotrophic bacterial abundance and production presented clear differences between MAW and Mediterranean water, with higher values at those stations under the influence of the Algerian Current. In general, greater differences were observed in production than in biomass variables. The photosynthetic parameters (derived from P–E relationships) and integrated primary production (range 189–645 mg m⁻² d⁻¹) responded greatly to the different hydrological conditions. The mesoscale phenomena inducing fertilization caused a 2 to 3-fold increase in primary production rates. The relatively high values found within the cyclonic eddy suggest that, although short-lived in comparison with anticyclonic eddies, these eddies may produce episodic increases of biological production not accounted for in previous surveys in the region.     

Biogeochemical responses to late-winter storms in the Sargasso Sea,II: Increased rates of biogenic silica production and export

Previous studies measuring biogenic silica production in the Sargasso Sea, all conducted when no phytoplankton bloom was in progress, have reported a mean rate of 0.4 mmol Si m^?2 d^?1 and maximum rate of 0.9 mmol Si m^?2 d^?1, the lowest rates yet recorded in any ocean habitat. During February/March of 2004 and 2005 we studied the effects of late-winter storms prior to seasonal stratification on the production rate, standing stock and vertical export of biogenic silica in the Sargasso Sea. In 2004, alternating storm and stratification events provided pulsed input of nutrients to the euphotic zone. In contrast, nearly constant storm conditions in 2005 caused the mixed layer to deepen to ～350 m toward the end of the cruise. Biogenic silica production rates in the upper 140 m were statistically indistinguishable between years, averaging ～1.0 mmol Si m^?2 d^?1. In early March 2004, a storm event entrained nutrients into the euphotic zone and, upon stabilization, vertically integrated biogenic silica in the upper 140 m nearly doubled in 2 days. Within 4 days, 75–100% of the accumulated biogenic silica was exported, sustaining a flux to 200 m of ～0.5 mmol Si m^?2 d^?1 (4× greater than export measured during February and March in the mid-1990s). In 2005, destabilization without stratification increased biogenic silica flux at 200 m up to two-fold above previously measured export in late winter, with little or no increase in water-column biogenic silica. Despite comprising <5% of total chlorophyll, diatoms accounted for an estimated 25–50% of the nitrate uptake in the upper 140 m and 35–97% of the particulate organic nitrogen export from the upper 200 m during both cruise periods. These previously unobserved brief episodes of diatom production and export in response to late-winter storms increase the estimated production and export of diatom-derived material in the Sargasso Sea in late winter by >150%, and increase estimated annual biogenic silica production in this region by ～8%.     

Occurrence and turnover of DMSP and DMS in deep waters of the Ross Sea,Antarctica

High concentrations of the phytoplankton metabolite dimethylsulfoniopropionate (DMSP) and its degradation product dimethylsulfide (DMS) are associated with blooms of Phaeocystis antarctica in the Ross Sea, Antarctica. Episodic and rapid vertical export of Phaeocystis biomass to deep water has been reported for the Ross Sea, therefore we examined the distribution and microbial consumption rates of DMSP and DMS throughout the sub-euphotic water column. Total DMSP (dissolved+particulate; DMSPt) was present at 0.5–22 nM at depths between 70 and 690 m during both the early bloom (November) and the late bloom (January). Sub-euphotic peaks of DMSP were sometimes associated with mid-water temperature maxima, and elevated DMSP below 70 m was found mainly in water masses characterized as Modified Circumpolar Deep Water or Antarctic Shelf Water. Overall, 50–94% of the integrated water-column DMSPt was found below the euphotic zone. At one station during the early bloom, local maxima of DMSPt (14 nM) and DMS (20 nM) were observed between 113 and 240 m and these maxima corresponded with high chlorophyll a concentrations, P. antarctica cell numbers, and Fv/Fm (the quantum yield of photosystem II). During the late bloom, a sub-euphotic maximum of DMSPt (15.8 nM) at 250 m cooccurred with peaks of chlorophyll a concentration, DMSP lyase activity, bacterial production and dissolved DMSP consumption rates. DMSP turnover contributed ～12% of the bacterial carbon demand between 200 and 400 m. DMS concentrations peaked at 286 m but the maximum concentration (0.42 nM) was far lower than observed during the early bloom, probably because of relatively rapid biological consumption of DMS (1–3 turnovers per day) which, in turn, contributed to elevated dissolved dimethylsulfoxide (DMSO) concentrations. Relatively stable DMSPt distributions at some sites suggest that rapid sinking of Phaeocystis biomass is probably not the major mechanism responsible for mesopelagic DMSP accumulations. Rather, subduction of near-surface water masses, lateral advective transport or trapping of slowly sinking P. antarctica biomass in intermediate water masses are more likely mechanisms. We found that a culture of P. antarctica maintained cellular integrity during 34 days of darkness, therefore the presence of intact cells (and DMSP) at depth can be explained even under a slow sinking/advection scenario. Whatever the mechanism, the large pools of DMSP and DMS below the euphotic zone suggest that export exerts a control on potential DMS emission from the surface waters of the Ross Sea.     

Reduced iron associated with secondary nitrite maxima in the Arabian Sea

Dissolved iron and Fe(II) were measured in the oxygen minimum zone (OMZ) of the Arabian Sea in September 2004. The OMZ is a well-demarcated feature characterized by high rates of denitrification, and a deep nitrite maximum coinciding with oxygen levels below 1 μmol L⁻¹. This zone is significantly enriched in dissolved Fe relative to overlying and underlying waters and up to 50% of the dissolved Fe is present as Fe(II). The maxima in Fe(II) are at the same depth as the deep nitrite maxima, centered around 200–250 m. They coincide with a local maximum in total dissolved Fe, suggesting that Fe accumulates at this depth because of the greater solubility of Fe(II) over Fe(III). Fe(II) is thermodynamically unstable even at submicromolar oxygen levels, so active biological reduction is the most plausible source. To our knowledge, this is the first report of a potential link between Fe reduction, elevated dissolved Fe concentrations, and nitrite accumulation within an OMZ. Denitrification has a high Fe requirement associated with the metalloenzymes for nitrate and nitrite reduction, so in situ redox cycling of Fe has important implications for the nitrogen cycle.     

Diurnal changes of photosynthetic quantum yield in the intertidal macroalga Sargassum thunbergii under simulated tidal emersion conditions

In this study, a three-way factorial experimental design was used to investigate the diurnal changes of photosynthetic activity of the intertidal macroalga Sargassum thunbergii in response to temperature, tidal pattern and desiccation during a simulated diurnal light cycle. The maximum (F_v/F_m) and effective (Φ_PSII) quantum yields of photosystem II (PSII) were estimated by chlorophyll fluorescence using a pulse amplitude modulated fluorometer. Results showed that this species exhibited sun-adapted characteristics, as evidenced by the daily variation of F_v/F_m and Φ_PSII. Both yield values decreased with increasing irradiance towards noon and recovered rapidly in the afternoon suggesting a dynamic photoinhibition. The photosynthetic quantum yield of S. thunbergii thalli varied significantly with temperature, tidal pattern and desiccation. Thalli were more susceptible to light-induced damage at high temperature of 25 °C and showed complete recovery of photosynthetic activity only when exposed to 8 °C. In contrast with the mid-morning low tide period, although there was an initial increase in photosynthetic yield during emersion, thalli showed a greater degree of decline at the end of emersion and remained less able to recover when low tide occurred at mid-afternoon. Short-term air exposure of 2 h did not significantly influence the photosynthesis. However, when exposed to moderate conditions (4 h desiccation at 15 °C or 6 h desiccation at 8 °C), a significant inhibition of photosynthesis was followed by partial or complete recovery upon re-immersion in late afternoon. Only extreme conditions (4 h desiccation at 25 °C or 6 h desiccation at 15 °C or 25 °C) resulted in the complete inhibition, with little indication of recovery until the following morning, implying the occurrence of chronic PSII damage. Based on the magnitude of effect, desiccation was the predominant negative factor affecting the photosynthesis under the simulated daytime irradiance period. These results may explain the distribution pattern of this species in natural habitats, where it is generally restricted to tide pools in the intertidal zone of wave-swept rocky shores which could provide shelter from desiccation stress during low tide.     

Seasonal and spatial variability in plankton production and respiration in the Subtropical Gyres of the Atlantic Ocean

Euphotic zone plankton production (P) and respiration (R) were determined from the in vitro flux of dissolved oxygen during six latitudinal transects of the Atlantic Ocean, as part of the Atlantic Meridional Transect (AMT) programme. The transects traversed the North and South Atlantic Subtropical Gyres (N gyre, 18–38°N; S gyre, 11–35°S) in April–June and September–November 2003–2005. The route and timing of the cruises enabled the assessment of the seasonal variability of P, R and P/R in the N and S gyres, and the comparison of the previously unsampled N gyre centre with the more frequently sampled eastern edge of the gyre. Mean euphotic zone integrated rates (±SE) were P=63±23 (n=31), R=69±22 (n=30) mmol O₂ m⁻² d⁻¹ in the N gyre; and P=58±26 (n=30), R=62±24 (n=30) mmol O₂ m⁻² d⁻¹ in the S gyre. Overall, the N gyre was heterotrophic (R>P) and it was more heterotrophic than the S gyre, but the metabolic balance of both gyres changed with season. Both gyres were net heterotrophic in autumn, and balanced in spring. This seasonal contrast was most pronounced for the S gyre, because it was more autotrophic than the N gyre during spring. This may have arisen from differences in nitrate availability, because spring sampling in the S gyre coincided with periods of deep mixing to the nitracline, more frequently than spring sampling within the N gyre. Our results indicate that the N gyre is less heterotrophic than previous estimates suggested, and that there is an apparent decrease in R from the eastern edge to the centre of the N gyre, possibly indicative of an allochthonous organic carbon source to the east of the gyre.     

Dissolved organic carbon and apparent oxygen utilization in the Atlantic Ocean

Dissolved organic carbon (DOC) distributions along two Atlantic Meridional Transects conducted in 2005 in the region between 47°N and 34°S showed clear latitudinal patterns. The DOC concentrations in the epipelagic zone (0–100 m) were the highest (70–90 µM) in tropical and subtropical waters with stable mixed layers, and lowest (50–55 µM) at the poleward extremities of the transects due to deep convective mixing supplying low DOC waters to the surface. A decrease in DOC occurred with depth, and lowest DOC concentrations (41–45 µM) in the 100–300 m depth range were observed in the equatorial region due to upwelling of low DOC waters. A strong relationship between DOC and AOU was observed in the σ–t 26–26.5 isopycnal layer which underlies the euphotic zone and outcrops at the poleward extremities of the North and South Atlantic Subtropical Gyres (NASG and SASG) in the region ventilating the thermocline waters. Our observations reveal significant north–south variability in the DOC–AOU relationship. The gradient of the relationship suggests that 52% of the AOU in the σ–t 26–26.5 density range was driven by DOC degradation in the NASG and 36% in the SASG, with the remainder due to the remineralisation of sinking particulate material. We assess possible causes for the greater contribution of DOC remineralisation in the NASG compared to the SASG.     

Biogeochemical responses to late-winter storms in the Sargasso Sea,I—Pulses of primary and new production

We have hypothesized that the weekly/biweekly passage of winter storms in the subtropical open ocean destabilizes the water column leading to pulsed NO₃^? inputs, resulting in new production that is not accounted for in most annual estimates. This paper presents data on nitrogen and carbon cycling in the Sargasso Sea at approximately daily resolution, during the period prior to seasonal stratification in 2004 and 2005; these data permit us to assess the importance of winter storms for introducing NO₃^? and the contribution of these inputs to annual new and export production. The two sampling years were in stark contrast to each other with 2004 characterized by periods of relative calm between winter storms, and 2005 characterized by nearly continuous storm activity. As a result, temporal variability in mixed layer depth (MLD) and euphotic zone [NO₃^?] were very different between years. MLDs in 2004 increased to >150 m in response to the passage of storms and then rapidly shoaled to <100 m leading to the pulsed injection of NO₃^? (～100 nmol l^?1) into the lower half of the euphotic zone, while in 2005 MLDs were consistently >300 m and euphotic zone [NO₃^?]>100 nmol l^?1. Despite the very different [NO₃^?], rates of daily NO₃^? uptake were similar from year to year because of significant nocturnal uptake in 2004. Similar rates of new production did not translate into similar rates of particulate nitrogen and carbon export however, as observed export from the upper 200 m was 2–5-fold greater in 2004 than in 2005. Furthermore, the decrease of particulate nitrogen and carbon flux with depth between 200 and 400 m in 2004 was substantially lower than in 2005; this is consistent with the observed biological response in which diatoms and coccolithophores exhibited rapid growth following pulsed NO₃^? inputs in 2004. A combination of data from the Bermuda Testbed Mooring, which provides a longer temporal record than the cruise, and the observations presented in this study show that in the winter of 2004, there were 8–10 storm events that likely resulted in pulsed NO₃^? inputs. Summed over all the events, new production prior to seasonal stratification was estimated to be ～0.12–0.18 mol N m^?2 or ～14–21% of current annual estimates.