Depth-integrated Production Research Articles

Cyanobacteria net community production in the Baltic Sea as inferred from profiling <i>p</i>CO<sub>2</sub> measurements

Abstract. Organic matter production by cyanobacteria blooms is a major environmental concern for the Baltic Sea, as it promotes the spread of anoxic zones. Partial pressure of carbon dioxide (pCO2) measurements carried out on Ships of Opportunity (SOOP) since 2003 have proven to be a powerful tool to resolve the carbon dynamics of the blooms in space and time. However, SOOP measurements lack the possibility to directly constrain depth-integrated net community production (NCP) in moles of carbon per surface area due to their restriction to the sea surface. This study tackles the knowledge gap through (1) providing an NCP best guess for an individual cyanobacteria bloom based on repeated profiling measurements of pCO2 and (2) establishing an algorithm to accurately reconstruct depth-integrated NCP from surface pCO2 observations in combination with modelled temperature profiles. Goal (1) was achieved by deploying state-of-the-art sensor technology from a small-scale sailing vessel. The low-cost and flexible platform enabled observations covering an entire bloom event that occurred in July–August 2018 in the Eastern Gotland Sea. For the biogeochemical interpretation, recorded pCO2 profiles were converted to CT*, which is the dissolved inorganic carbon concentration normalised to alkalinity. We found that the investigated bloom event was dominated by Nodularia and had many biogeochemical characteristics in common with blooms in previous years. In particular, it lasted for about 3 weeks, caused a CT* drawdown of 90 µmol kg−1, and was accompanied by a sea surface temperature increase of 10 ∘C. The novel finding of this study is the vertical extension of the CT* drawdown up to the compensation depth located at around 12 m. Integration of the CT* drawdown across this depth and correction for vertical fluxes leads to an NCP best guess of ∼1.2 mol m−2 over the productive period. Addressing goal (2), we combined modelled hydrographical profiles with surface pCO2 observations recorded by SOOP Finnmaid within the study area. Introducing the temperature penetration depth (TPD) as a new parameter to integrate SOOP observations across depth, we achieve an NCP reconstruction that agrees to the best guess within 10 %, which is considerably better than the reconstruction based on a classical mixed-layer depth constraint. Applying the TPD approach to almost 2 decades of surface pCO2 observations available for the Baltic Sea bears the potential to provide new insights into the control and long-term trends of cyanobacteria NCP. This understanding is key for an effective design and monitoring of conservation measures aiming at a Good Environmental Status of the Baltic Sea.

Dynamics of primary productivity in relation to submerged vegetation of a shallow, eutrophic lagoon: A field and mesocosm study.

Aquatic ecosystems nowadays are under constant pressure, either from recent or historical events. In most systems with increased nutrient supply, submerged macrophytes got replaced by another stable state, dominated by phytoplankton as main primary producer. Yet, reducing the nutrient supply did not yield the aimed goal of restored habitats for submerged macrophytes in systems worldwide. The question arises, why submerged macrophytes do not re-colonize, and if they are actually competitive. Therefore, primary production assays were conducted in ex-situ bentho-pelagic mesocosms and compared to the actual ecosystem, a turbid brackish lagoon of the southern Baltic Sea. Mesocosm were either manipulated to be colonized by macrophytes, or stayed phytoplankton dominated. Oxygen evolution was monitored over a period of five months in 5 min (mesocosms) to 10 min (ecosystem) intervals. Surface and depth-integrated production was calculated to analyse seasonal and areal resolved production patterns. It was found that macrophyte mesocosms were more stable, when considering only surface O2 production. However, calculating depth-integrated production resulted in net-heterotrophy in both shallow mesocosms approaches and the actual ecosystem. This heterotrophy is likely mediated by sediment respiration and POC accumulation in mesocosms, and a low share of productive to respiring water column in the actual ecosystem. Therefore, it seems unlikely that macrophytes will re-settle, as constant net-heterotrophy may allow for high-nutrient turnover at sediment-water interfaces and within the water column, favouring phytoplankton. These results will assist decision makers in developing more effective restoration measures that can mitigate the negative effects of eutrophication on ecosystem function and services.

Synoptic Spatio-Temporal Variability of the Photosynthetic Productivity of Microphytobenthos and Phytoplankton in a Tidal Estuary

Tidal estuaries are regarded as highly important ecosystems, mostly due to their high primary productivity and associated role as carbon sinks. In these ecosystems, primary productivity is mainly due to the photosynthetic carbon fixation by phytoplankton and microphytobenthos. The productivity of the two communities has been mostly studied separately, and directly comparable estimates of their carbon fixation rates in the same estuary are relatively scarce. The present study aimed to characterize the spatio-temporal variability of the productivity of phytoplankton and microphytobenthos in a tidal estuary, the Ria de Aveiro (Portugal), and to estimate the annual ecosystem-level budget for carbon fixation by the two communities. Productivity rates were determined based on synoptic in situ measurements of absolute rates of electron transport rate of photosystem II, using PAM fluorometry. Chlorophyll fluorescence indices were accompanied by measurements of salinity, temperature, water turbidity, solar irradiance, and planktonic and benthic microalgal biomass. Measurements were carried out hourly, along four spring-neap tidal cycles distributed along one year, on three sites of the estuary. The most pronounced trends in the spatio-temporal variability of the photophysiology and productivity of the two communities were the following: (i) maximum biomass and productivity were reached later for microphytobenthos (summer-autumn) than for phytoplankton (spring-summer); (ii) the absorption cross-section of PSII was generally higher for phytoplankton; (iii) the two groups showed a similar photoacclimation state, but microphytobenthos appeared as high light-acclimated when compared to phytoplankton. Biomass-specific productivity was on average higher for phytoplankton than for microphytobenthos, averaging 68.0 and 19.1 mg C mg Chl a-1 d-1, respectively. However, areal depth-integrated production rates were generally higher for the microphytobenthos than for the phytoplankton, averaging 264.5 and 140.0 mg C m-2 d-1, respectively. On an annual basis, phytoplankton productivity averaged 49.9 g C m-2 yr-1 while the productivity of microphytobenthos averaged 105.2 g C m-2 yr-1. When upscaling to the whole estuary, annual primary production rates of phytoplankton and microphytobenthos reached 4894.3 and 7534.0 t C yr-1, respectively, representing 39.4% and 60.6% of the combined total of 12428.3 t C yr-1 determined for the two communities in the Ria de Aveiro.

Patterns of the Kara Sea primary production in autumn: Biotic and abiotic forcing of subsurface layer

Primary production and fundamental environmental factors were measured during September–October 1993, 2007 and 2011 in the Kara Sea. Relationships between the depth-integrated primary production (PPint), the surface chlorophyll a (Chl0) concentration and the maximum chlorophyll specific carbon rate within water column (Pbopt) had shown that only 12% of PPint variability were determined by Chl0 and there were strong correlations between PPint and Pbopt (R2=0.64). Thus, in the autumn PPint values were largely influenced by the phytoplankton assimilation activity. At the end of a vegetative season high (close or above 1mgm−3) values of Chl0 were not an index of phytoplankton productivity within photosynthetic layer where the organic matter synthesis rate was low which testifies the lack of correlation between Chl0 and PPint (R2=0.12) and between Chl0 and depth-integrated chlorophyll a (chl a) (R2=0.22). This conclusion is based on the low assimilation activity within water column and small thickness of the photosynthetic layer. The latter corresponds to low insolation and water transparency. The lack of correlation between surface and depth-integrated productivity parameters is the challenge for satellite PPint estimations. In turn PPint and Pbopt depended mainly on photosynthetically available radiation (PAR) and weakly related to the nitrogen and phosphorus concentrations. At the end of a vegetative season PAR level, apparently, should be considered as the main factor for primary production in the Kara Sea. Comparison between the integrated primary production from shipboard in September and the simulated evaluations of productivity (PP models) suggests that PP models overestimate the in situ PPint by a factor of 3–7 in the different Kara Sea regions. Improving of Kara Sea primary production estimations implies the development of regional satellite chl a algorithm and local primary production model considering specific features of PP in this Arctic Ocean region.

Spectrally resolved efficiencies of carbon monoxide (CO) photoproduction in the western Canadian Arctic: particles versus solutes

Abstract. Spectrally resolved efficiency (i.e. apparent quantum yield, AQY) of carbon monoxide (CO) photoproduction is a useful indicator of substrate photoreactivity and a crucial parameter for modeling CO photoproduction rates in the water column. Recent evidence has suggested that CO photoproduction from particles in marine waters is significant compared to the well-known CO production from chromophoric dissolved organic matter (CDOM) photodegradation. Although CDOM-based CO AQY spectra have been extensively determined, little is known of this information on the particulate phase. Using water samples collected from the Mackenzie estuary, shelf, and Canada Basin in the southeastern Beaufort Sea, the present study for the first time quantified the AQY spectra of particle-based CO photoproduction and compared them with the concomitantly determined CDOM-based CO AQY spectra. CO AQYs of both particles and CDOM decreased with wavelength but the spectral shape of the particulate AQY was flatter in the visible regime. This feature resulted in a disproportionally higher visible light-driven CO production by particles, thereby increasing the ratio of particle- to CDOM-based CO photoproduction with depth in the euphotic zone. In terms of depth-integrated production in the euphotic zone, CO formation from CDOM was dominated by the ultraviolet (UV, 290–400 nm) radiation whereas UV and visible light played roughly equal roles in CO production from particles. Spatially, CO AQY of bulk particulate matter (i.e. the sum of organics and inorganics) augmented from the estuary and shelf to the basin while CO AQY of CDOM trended inversely. Water from the deep chlorophyll maximum layer revealed higher CO AQYs than did surface water for both particles and CDOM. CO AQY of bulk particulate matter exceeded that of CDOM on the shelf and in the basin, but the sequence reversed in the estuary. Without consideration of the potential role of metal oxides (e.g. iron oxides) in particle photochemistry, mineral absorption-corrected CO AQY of particulate organic matter (POM) could, however, surpass its CDOM counterpart in all three sub-regions and displayed magnitudes in the estuary that overtook those in shelf and offshore waters. In terms of CO photoproduction, POM was thus more photoreactive than CDOM, irrespective of the organic matter's origins (i.e. terrigenous or marine). Riverine CDOM exhibited higher photoreactivity than marine CDOM and land-derived POM appeared more photoreactive than marine POM. AQY-based modeling indicates that CO photoproduction in the study area is underestimated by 12–32% if the particulate term is ignored.

Primary and new production in the thermocline ridge region of the southern Indian Ocean during the summer monsoon

New and primary production, measured using 15N and 13C tracers, integrated over the euphotic zone (mostly upper 80-100 m) varied from 7.5 to 14.1 mmol N m-2d-1(average 10.8 mmol N m-2d-1) and 2.7 to 17.7 mmol C m-2d-1 (average 11.3 mmol C m-2d-1), respectively, in the tropical India Ocean (TIO). Shoaling of thermocline in the thermocline ridge located in the tropical Indian Ocean (TR) influences new and total productivity by supplying nutrients to the upper layers. At the same time, N-limited conditions prevail to the north of the TR region reducing biological productivity. While higher surface light intensity reduces the surface productivity, it enhances the overall depth-integrated productivity. Productivity at different locations within the region is primarily controlled by the supply of N-nutrients from below. In contrast, at a given location with the available nutrient pool, productivity at different depths is determined by light availability. The region has potential for moderate export production, as do the other parts of the Indian Ocean; the maximum possible f-ratio is found to be 0.56, suggesting that it is capable of exporting a significant part of the total production to the deep, under favorable conditions (i.e., availability of N-nutrients). A comparison with earlier data suggests no significant temporal trend in the productivity here over the past three decades.

Vertical distribution of prokaryote production and abundance in the mesopelagic and bathypelagic layers of the Canada Basin, western Arctic: Implications for the mode and extent of organic carbon delivery

The vertical distributions of prokaryote heterotrophic production (3H-leucine incorporation rate) and abundance were investigated in the meso- and bathy-pelagic layers of the Canada Basin, western Arctic Ocean, during September 2009. Prokaryote production and abundance were high in the Pacific-origin water mass located in the upper mesopelagic layer (depth, 100–200m). Below the halocline layer (depth, 300–3000m), both the production and abundance decreased with depth, with log–log regression slopes of −1.33 and −0.77, respectively. Depth-integrated production and biomass in the meso- and bathy-pelagic layers was three- to five-fold lower than the corresponding values reported in the subpolar regions, whereas they were close to or lower than the corresponding values in oligotrophic subtropical regions. Prokaryote turnover times were estimated to be 1.1 and 6.1 years for meso- and bathy-pelagic layers, respectively, with the latter being among the longest turnover times reported for oceanic basins. We estimated prokaryote carbon demand in the water column (100–3000m) to be on the order of 11mgCm−2d−1, which largely exceeds (by 38-fold) the sinking particulate organic carbon flux at depths of 120–200m reported in the literature. This large carbon imbalance may be partly explained by organic carbon delivery by lateral intrusion of the Pacific-origin water mass into the upper mesopelagic layer.

Short-Term and Interannual Variability in Primary Production in the Low-Salinity Zone of the San Francisco Estuary

We measured primary production during spring–summer 2006–2007 to determine the carbon supply to the low-salinity pelagic food web of the San Francisco Estuary (SFE). Weekly or biweekly samples were taken at three stations of fixed salinity for size-fractionated primary production and biomass, both as chlorophyll and from biovolume based on counts. Error variance in productivity estimates arose mainly from the depth integration of 14C uptake, showing the importance of productivity measurements at high light levels for estimates of depth-integrated production. Temporal and spatial variability in production were surprisingly small. Combining data from this study with long-term monitoring data, productivity and biomass were variable in time and salinity but without persistent patterns and with infrequent blooms. Production within the low-salinity zone was unresponsive to variation in freshwater flow, in contrast to findings in other estuaries where nutrient loading drives variability in production and other regions of the SFE where production responds to residence time or to stratification. Estimated annual primary production was only 25 and 31 g C m−2 year−1 during 2006 and 2007, only half of it in cells >5 μm. These results imply that phytoplankton provided poor food web support for higher trophic levels, probably contributing to the long-term decline in fish abundance in the brackish to freshwater region of the estuary.

Calculations of in-snow NO2and OH radical photochemical production and photolysis rates: A field and radiative-transfer study of the optical properties of Arctic (Ny-Ålesund, Svalbard) snow

[1] Depth-integrated production rates of OH radicals and NO2 molecules from snowpacks in Ny-Alesund, Svalbard, are calculated from fieldwork investigating the light penetration depth (e-folding depth) and nadir reflectivity of snowpacks during the unusually warm spring of 2006. Light penetration depths of 8.1, 11.3, 5.1, and 8.2 cm were measured for fresh, old, marine-influenced, and glacial snowpacks, respectively (wavelength 400 nm). Radiative-transfer calculations of the light penetration depths with reflectivity measurements produced scattering cross sections of 5.3, 9.5, 20, and 25.5 m2 kg−1 and absorption cross sections of 7.7, 1.4, 3.4, and 0.5 cm2 kg−1 for the fresh, old, marine-influenced, and glacial snowpacks, respectively (wavelength 400 nm). Photolysis rate coefficients, J, are presented as a function of snow depth and solar zenith angle for the four snowpacks for the photolysis of H2O2 and NO3−. Depth-integrated production rates of hydroxyl radicals are 1270, 2130, 950, and 1850 nmol m−2 h−1 (solar zenith angle of 60°) for fresh, old, marine-influenced, and glacial snowpacks, respectively. Depth-integrated production rates of NO2 are 32, 56, 11, and 22 nmol m−2 h−1 (solar zenith angle of 60°) for the fresh, old, marine-influenced, and glacial snowpacks, respectively. The uncertainty of repeated light penetration depth measurement was determined to be ∼20%, which propagates into a 20% error in depth-integrated production rates. A very simple steady state hydroxyl radical calculation demonstrates that a pseudo first-order loss rate of OH radicals of ∼102–104 s−1 is required in snowpack. The snowpacks around Ny-Alesund are thick enough to be considered optically infinite.

The importance of considering depth-resolved photochemistry in snow: a radiative-transfer study of NO2 and OH production in Ny-Ålesund (Svalbard) snowpacks

AbstractSolar visible radiation can penetrate 2–30 cm (e-folding depth) into snowpacks and photolyse nitrate anions and hydrogen peroxide contained in the snow. Photolysis rate coefficients, J, for NO3− and H2O2 photolysis are presented for a melting and a fresh snowpack at Ny-Ålesund, Svalbard. Calculations of (a) transfer velocities, υ, and molecular fluxes of gaseous NO2 from the snowpack and (b) depth-integrated production rates of OH radicals within the snowpack are presented. The results show the importance of considering the depth dependence, i.e. not just the snow surface, when modelling snowpack photochemistry. Neglecting photochemistry under the snow surface can result in an apparent larger molecular flux of NO2 from NO3− photolysis than the melting snowpack. However, when the depth-resolved molecular fluxes of NO2 within the snowpack are calculated, a larger NO2 flux may be apparent in the melting snowpack than the fresh snowpack. For solar zenith angles of 60°, 70° and 80° the modelled molecular fluxes of NO2 from fresh snowpack are 11.6, 5.6 and 1.7 nmol m−2 h−1, respectively, and those for melting snowpack are 19.7, 9.1 and 2.9 nmol m−2 h−1, respectively.

Hydroxyl (OH) radical production rates in snowpacks from photolysis of hydrogen peroxide (H 2O 2) and nitrate (NO 3−)

Atmospheric chemistry directly above snowpacks is strongly influenced by ultraviolet (UV) radiation initiated emissions of chemicals from the snowpack. The emission of gases from the snowpack to the atmosphere is in part due to chemical reactions between hydroxyl radical, OH (produced from photolysis of hydrogen peroxide (H 2O 2) or nitrate (NO 3 −)) and impurities in the snowpack. The work presented here is a radiative-transfer modelling study to calculate the depth-integrated production rates of hydroxyl radical from the photolysis of hydrogen peroxide and nitrate anion in snow for four different snowpacks and for solar zenith angles 30°–90°. This work also demonstrates the importance of hydrogen peroxide photolysis to produce hydroxyl radical relative to nitrate photolysis with (a) different snowpacks, (b) different ozone column depths, and (c) snowpack depths. The importance of hydrogen peroxide photolysis over nitrate photolysis for hydroxyl radical production increases with increasing depth in snowpack, column ozone depth, and solar zenith angle. With a solar zenith angle of 60° the production of hydroxyl radical from hydrogen peroxide photolysis accounts for 91–99% of all hydroxyl radical production from hydrogen peroxide and nitrate photolysis.

Phytoplankton as an increasingly compound photosynthetic system: an historical perspective

The original suggestion that phytoplankton can be viewed and modelled as a compound photosynthetic system combined light gradients with depth and biological plus chemical homogeneity. Subsequent experience often involved biological and chemical differentiation in depth and time. Such differentiation is surveyed; it can add to the compound structure, and modify the kinetics of depth-integrated photosynthetic production. Further compounding possibilities are introduced by alternatives in carbon input, output and storage; also, conceptually, by the level of approach adopted. These features are discussed in historical perspective.

Seasonal variations in production and consumption rates of dissolved organic carbon in an organic-rich coastal sediment

Dissolved organic carbon (DOC) concentrations in anoxic marine sediments are controlled by at least three processes: 1. (1) production of nonvolatile dissolved compounds, such as peptides and amino acids, soluble saccharides and fatty acids, via hydrolysis of particulate organic carbon (POC). 2. (2) conversion of these compounds to volatile fatty acids and alcohols by fermentative bacteria. 3. (3) consumption of volatile fatty acids and alcohols by terminal bacteria, such as sulfate reducers and methanogens. We monitored seasonal changes in concentration profiles of total DOC, nonacid-volatile (NAV) DOC and acid-volatile (AV) DOC in anoxic sediment from Cape Lookout Bight, North Carolina, USA, in order to investigate the factors that control seasonal variations in rates of hydrolysis, fermentation, and terminal metabolism. During the winter months, DOC concentrations increased continuously from 0.2 mM in the bottomwater to ~4 mM at a depth of 36 cm in the sediment column. During the summer, a large DOC maximum developed between 5 and 20 cm, with peak concentrations approaching 10 mM. The mid-depth summertime maximum was driven by increases in both NAV- and AV-DOC concentrations. Net NAV-DOC reaction rates were estimated by a diagenetic model applied to NAV-DOC concentration profiles. Depth-integrated production rates of NAV-DOC increased from February through July, suggesting that net rates of POC hydrolysis during this period are controlled by temperature. Net consumption of NAV-DOC during the late summer and early fall suggests reduced gross NAV-DOC production rates, presumably due to a decline in the availability of labile POC. A distinct subsurface peak in AV-DOC concentration developed during the late spring, when the sulfate depletion depth shoaled from 25 to 10 cm. We hypothesize that the AV-DOC maximum results from a decline in consumption by sulfate-reducing bacteria (due to sulfate limitation) and a lag in the development of an active population of methanogenic bacteria. A diagenetic model that incorporates a lag period in the sulfate reducer-methanogen transition successfully simulates the timing, magnitude, depth and shape of the AV-DOC peak.

A numerical study of primary production related to vertical turbulent diffusion with special reference to vertical motions of the phytoplankton cells in nutrient and light fields

Assuming stationary physical processes, in particular the light field and turbulent activity [ K( z)], we described steady-state and convergent solutions obtained from a simple time-dependent vertical model of phytoplankton dynamics. Simulations included vertical turbulent motions experienced by the cells in the light and nutrient fields. Parallel simulations made with a classical formulation of phytoplankton growth, i.e., neglecting vertical turbulent motions, are discussed. From two typical situations of stratification in the Western Mediterranean, we identified two distinct systems of new production, as the consequence of Low (LTR) and High Turbulent Regime (HTR) in the photic zone respectively. Data from the Prolig-II (1985) and Almofront-I (1991) cruises supported the LTR system of new production. The results of the second part of the Mediprod-I (1969) cruise show several patterns that specifically appeared in the HTR simulation. Regenerated production was not influenced by the turbulent activity situation. In natural conditions, regenerated production depends on the specific phytoplankton-grazers system that develops according to the level of new production; such ecological dynamics were not considered in our model. Differences with the reference model changed the relationships between the vertical distributions of biomass and new production. Particularly, the HTR simulation led to distinct vertical distribution of biomass and new production. Such a pattern did not occur with the reference model. Although the vertical turbulent motions affected both the level and vertical distribution of new production, a significant effect on the depth-integrated production finally depends on how the phytoplankton biomass interacts with its environmental conditions. It is shown that the minimum of K( z) in the euphotic zone determined the system of new production, whereas its values below the euphotic zone scaled the production and biomass levels. The two distinct systems of new production, LTR and HTR, may be diagnosed from simple cast measurement by examining the relationships between the distributions of parameters implicated in new production (biomass maximum, nitracline and oxycline) and the density profile.