New constraints on biological production and mixing processes in the South China Sea from triple isotope composition of dissolved oxygen

This research has been supported by the Ministry of Science and Technology, Taiwan (grant no. 108-2111-M-001-011-MY3) and the Academia Sinica (grant no. AS-IA-109-M03).The South China Sea (SCS) is the world's largest marginalsea, playing an important role in the regional biogeochemical cycling ofcarbon and oxygen. However, its overall metabolic balance, primaryproduction rates and links to East Asian Monsoon forcing remain poorlyconstrained. Here, we report seasonal variations in triple oxygen isotopecomposition (17Δ) of dissolved O2, a tracer for biologicalO2, gross primary production (GP; inferred from δ17O andδ18O values) and net community production (NP; evaluated fromoxygen–argon ratios) from the SouthEast Asian Time-series Study (SEATS) inthe SCS. Our results suggest rather stable mixed-layer mean GP rates of∼ 1500 ± 350 mg C m−2 d−1 and mean NP of∼ −13 ± 20 mg C m−2 d−1 during the summersouthwest monsoon season. These values indicate, within uncertainties andvariabilities observed, that the metabolism of the system was in netbalance. During months influenced by the stronger northeast monsoon forcing,the system appears to be more dynamic and with variable production rates,which may shift the metabolism to net autotrophy (with NP rates up to∼ 140 mg C m−2 d−1). Furthermore, our data from thedeeper regions show that the SCS circulation is strongly affected by monsoonwind forcing, with a larger part of the water column down to at least 400 mdepth fully exchanged during a winter, suggesting the 17Δ ofdeep O2 as a valuable novel tracer for probing mixing processes.Altogether, our findings underscore the importance of monsoon intensity onshifting the carbon balance in this warm oligotrophic sea and on drivingthe regional circulation pattern.Publisher PDFPeer reviewe

Activity and Distribution of Attached Bacteria in Chesapeake Bay

The purpose of this study was to further our understanding of the role of particle-associated bacteria in phytoplankton degradation in the Chesapeake Bay, USA, and to identify environmental parameters that control production by free and particle-associated bacteria. Surface and bottom waters at 10 stations along the length of the Bay were sampled over a 2 yr period. Samples were analyzed for temperature, salinity, chlorophyll, phaeophytin, particulate protein, thymidine incorporation (an estimate of bacterial growth rate), and bacterial total direct counts. Results demonstrated that freeliving bacteria were responsible for most of the total bacterial production, which was correlated with temperature and particulate protein, but not with chlorophyll. In contrast, attached bacteria were much more active than free-living bacteria on a per-cell basis. Cell-specific thymidine incorporation by freeliving bacteria correlated with temperature and particulate protein quantity, whereas that by attached bacteria correlated with temperature, particulate protein quantity and particulate organic quality, as indicated by extent of phytoplankton degradation

Variations in triple isotope composition of dissolved oxygen and primary production in a subtropical reservoir

The work was supported in part by MOST grants 101-2628-M-001-001-MY4 and 105-2111-M-001-006-MY3 to Academia Sinica.Lakes and reservoirs play an important role in the carbon cycle, and therefore monitoring their metabolic rates is essential. The triple oxygen-isotope anomaly of dissolved O2 [17Δ =  ln(1+δ17O) − 0.518  ×  ln(1 + δ18O)] offers a new, in situ, perspective on primary production, yet little is known about 17Δ from freshwater systems. We investigated the 17Δ together with the oxygen : argon ratio [Δ(O2 ∕ Ar)] in the subtropical Feitsui Reservoir in Taiwan from June 2014 to July 2015. Here, we present the seasonal variations in 17Δ, GP (gross production), NP (net production) and the NP ∕ GP (net to gross ratio) in association with environmental parameters. The 17Δ varied with depth and season, with values ranging between 26 and 205 per meg. The GP rates were observed to be higher (702 ± 107 mg C m−2 d−1) in winter than those (303 ± 66 mg C m−2 d−1) recorded during the summer. The overall averaged GP was 220 g C m−2 yr−1 and NP was −3 g C m−2 yr−1, implying the reservoir was net heterotrophic on an annual basis. This is due to negative NP rates from October to February (−198 ± 78 mg C m−2 d−1). Comparisons between GP rates obtained from the isotope mass balance approach and 14C bottle incubation method (14C–GP) showed consistent values on the same order of magnitude with a GP ∕ 14C–GP ratio of 1.2 ± 1.1. Finally we noted that, although typhoon occurrences were scarce, higher than average 17Δ values and GP rates were recorded after typhoon events.Publisher PDFPeer reviewe

The Effects of Light and Nitrate Levels on the Relationship Between Nitrate Reductase Activity and (NO3-)-N-15 Uptake: Field Observations in the East China Sea

Nitrate reductase activity (NRA) and 15NO3- uptake (NU) were determined in the East China Sea and the adjoining Kuroshio in May 1996, at six stations covering a range of hydrographic conditions: the nutrient-rich and fresher plume of Changjiang Diluted Water along the Chinese coast, the nutrient-rich upwelling Kuroshio Subsurface Water at the shelf edge northeast of Taiwan, the oligotrophic Kuroshio Surface Water and the mixing zones among these water masses on the shelf. The values of NRA in the surface mixed layer ranged between 16 and 0.1 nM-N h-1, whereas those of NU ranged between 37 and 1 nM-N h-1. Higher NRA and NU were found in the frontal zone between the coastal and shelf waters and in the upwelling zone, whereas the lowest values were found in the surface Kuroshio. The NRA/Chl a ratio increased linearly with increasing NU/primary production ratio in the sequence: Kuroshio \u3c coastal plume \u3c upwelling zone and mixing zones in the shelf. This is probably a reflection of the varying nutrient condition and the relative importance of NU in sustaining the biomass in these regions. In nitrate- and light-replete waters, the average NU/NRA was 1.0 +/- 0.3. NRA was linearly related to NU so that NU = 1.08 (+/- 0.07)NRA (r(2) = 0.79). Thus, NRA may be used for estimating NU in these waters. In nitrate deficient and light-replete waters, the average NU/NRA was 4 +/- 4. These high and variable values of NU/NRA might have been caused by an over-estimation of NU as a result of the stimulatory effect of the added 15NO3-N on phytoplankton growth. Thus, NRA may be a more reliable indicator of the rate of NO3- uptake in oligotrophic waters. In nitrate-replete and light-deficient waters, NU did not correlate well with NRA. The average NU/NRA was 0.7 +/- 0.7. These low and variable values of NU/NRA suggest a possible decoupling between NRA and NU. By using the relationship between NU and NRA in nitrate- and light-replete waters and the depth-integrated inventory of NRA in the photic zone at each station, NU in oligotrophic waters, the coastal plume, upwelling waters and shelf waters can be estimated to be 0.45, 1.55, 3.12, and 3.59 mg-N m-2 h-1 respectively. These values fall well within the range of previously reported values in similar types of water

Nitrate Anomaly in the Upper Nutricline in the Northern South China Sea - Evidence for Nitrogen Fixation

[1] Up to 2 μM of nitrate anomaly, N*, were found in the upper nutricline at the South East Asia Time-series Study (SEATS) site in the northern South China Sea (SCS). These concentrations were among the higher values reported in the Pacific and indicate the significant contribution of the remineralization of nitrogen-rich organic matter formed by nitrogen fixation to the nutrient dynamics of the area. The concentrations were systematically higher, by up to 2.5 μM, in the Fall through the early Spring, during the northeast monsoon, than in the Summer, suggesting that the impact of nitrogen fixation was higher during the former time period. This pattern is in phase with that of the atmospheric deposition of Asian dust to the northern SCS. The coherence is consistent with a coupling between nitrogen fixation and the availability of atmospherically derived iron

Variations in triple isotope composition of dissolved oxygen and primary production in a subtropical reservoir

Lakes and reservoirs play an important role in the carbon cycle, and therefore monitoring their metabolic rates is essential. The triple oxygen-isotope anomaly of dissolved O2 [17Δ =  ln(1+δ17O) − 0.518  ×  ln(1 + δ18O)] offers a new, in situ, perspective on primary production, yet little is known about 17Δ from freshwater systems. We investigated the 17Δ together with the oxygen : argon ratio [Δ(O2 ∕ Ar)] in the subtropical Feitsui Reservoir in Taiwan from June 2014 to July 2015. Here, we present the seasonal variations in 17Δ, GP (gross production), NP (net production) and the NP ∕ GP (net to gross ratio) in association with environmental parameters. The 17Δ varied with depth and season, with values ranging between 26 and 205 per meg. The GP rates were observed to be higher (702 ± 107 mg C m−2 d−1) in winter than those (303 ± 66 mg C m−2 d−1) recorded during the summer. The overall averaged GP was 220 g C m−2 yr−1 and NP was −3 g C m−2 yr−1, implying the reservoir was net heterotrophic on an annual basis. This is due to negative NP rates from October to February (−198 ± 78 mg C m−2 d−1). Comparisons between GP rates obtained from the isotope mass balance approach and 14C bottle incubation method (14C–GP) showed consistent values on the same order of magnitude with a GP ∕ 14C–GP ratio of 1.2 ± 1.1. Finally we noted that, although typhoon occurrences were scarce, higher than average 17Δ values and GP rates were recorded after typhoon events

Community production modulates coral reef pH and the sensitivity of ecosystem calcification to ocean acidification

Author Posting. © American Geophysical Union, 2017. This article is posted here by permission of American Geophysical Union for personal use, not for redistribution. The definitive version was published in Journal of Geophysical Research: Oceans 122 (2017): 745–761, doi:10.1002/2016JC012326.Coral reefs are built of calcium carbonate (CaCO3) produced biogenically by a diversity of calcifying plants, animals, and microbes. As the ocean warms and acidifies, there is mounting concern that declining calcification rates could shift coral reef CaCO3 budgets from net accretion to net dissolution. We quantified net ecosystem calcification (NEC) and production (NEP) on Dongsha Atoll, northern South China Sea, over a 2 week period that included a transient bleaching event. Peak daytime pH on the wide, shallow reef flat during the nonbleaching period was ∼8.5, significantly elevated above that of the surrounding open ocean (∼8.0–8.1) as a consequence of daytime NEP (up to 112 mmol C m−2 h−1). Diurnal-averaged NEC was 390 ± 90 mmol CaCO3 m−2 d−1, higher than any other coral reef studied to date despite comparable calcifier cover (25%) and relatively high fleshy algal cover (19%). Coral bleaching linked to elevated temperatures significantly reduced daytime NEP by 29 mmol C m−2 h−1. pH on the reef flat declined by 0.2 units, causing a 40% reduction in NEC in the absence of pH changes in the surrounding open ocean. Our findings highlight the interactive relationship between carbonate chemistry of coral reef ecosystems and ecosystem production and calcification rates, which are in turn impacted by ocean warming. As open-ocean waters bathing coral reefs warm and acidify over the 21st century, the health and composition of reef benthic communities will play a major role in determining on-reef conditions that will in turn dictate the ecosystem response to climate change.NSF Grant Number: 12205292017-07-3

Prokaryotic assemblages and metagenomes in pelagic zones of the South China Sea

Background: Prokaryotic microbes, the most abundant organisms in the ocean, are remarkably diverse. Despite numerous studies of marine prokaryotes, the zonation of their communities in pelagic zones has been poorly delineated. By exploiting the persistent stratification of the South China Sea (SCS), we performed a 2-year, large spatial scale (10, 100, 1000, and 3000 m) survey, which included a pilot study in 2006 and comprehensive sampling in 2007, to investigate the biological zonation of bacteria and archaea using 16S rRNA tag and shotgun metagenome sequencing. Results: Alphaproteobacteria dominated the bacterial community in the surface SCS, where the abundance of Betaproteobacteria was seemingly associated with climatic activity. Gammaproteobacteria thrived in the deep SCS, where a noticeable amount of Cyanobacteria were also detected. Marine Groups II and III Euryarchaeota were predominant in the archaeal communities in the surface and deep SCS, respectively. Bacterial diversity was higher than archaeal diversity at all sampling depths in the SCS, and peaked at mid-depths, agreeing with the diversity pattern found in global water columns. Metagenomic analysis not only showed differential %GC values and genome sizes between the surface and deep SCS, but also demonstrated depth-dependent metabolic potentials, such as cobalamin biosynthesis at 10 m, osmoregulation at 100 m, signal transduction at 1000 m, and plasmid and phage replication at 3000 m. When compared with other oceans, urease at 10 m and both exonuclease and permease at 3000 m were more abundant in the SCS. Finally, enriched genes associated with nutrient assimilation in the sea surface and transposase in the deep-sea metagenomes exemplified the functional zonation in global oceans. Conclusions: Prokaryotic communities in the SCS stratified with depth, with maximal bacterial diversity at mid-depth, in accordance with global water columns. The SCS had functional zonation among depths and endemically enriched metabolic potentials at the study site, in contrast to other oceans