Joint Effect of Freshwater Plume and Coastal Upwelling on Phytoplankton Growth Off the Changjiang River
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Biogeosciences, 11, 409–423, 2014 Open Access www.biogeosciences.net/11/409/2014/ doi:10.5194/bg-11-409-2014 Biogeosciences © Author(s) 2014. CC Attribution 3.0 License. Joint effect of freshwater plume and coastal upwelling on phytoplankton growth off the Changjiang River Y.-F. Tseng1, J. Lin2, M. Dai2, and S.-J. Kao1,2 1Research Center for Environmental Changes, Academia Sinica, Taipei, China 2State Key Laboratory of Marine Environmental Science, Xiamen University, Xiamen, China Correspondence to: S.-J. Kao ([email protected]) Received: 11 April 2013 – Published in Biogeosciences Discuss.: 26 June 2013 Revised: 6 December 2013 – Accepted: 12 December 2013 – Published: 28 January 2014 Abstract. The Changjiang (Yangtze) River discharges vast causing dramatic changes of nutrient status in adjacent es- amount of unbalanced nutrients (dissolved inorganic nitro- tuarine and coastal ecosystems (Jickells, 1998). These rivers gen and phosphorus with N / P ratio > 80 in general) into are well known to carry nutritional water with high ratios the East China Sea in summer. To study nutrient dynam- of dissolved inorganic nitrogen (DIN) to dissolved inorganic ics and P-stress potential for phytoplankton, a cruise was phosphorus (DIP), or N / P ratios. Such massive yet dispro- conducted in the Changjiang plume during summer 2011. portionate riverine nutrient input has caused an obvious “ex- With 3-D observations of nutrients, chlorophyll a (Chl a), cess N” (the concentration of DIN in excess of DIP) phe- and bulk alkaline phosphatase activity (APA), we concluded nomenon in nearby estuarine ecosystems (Galloway et al., that the Changjiang Diluted Water and coastal upwelling 2004). Ultimately, the continuing increase in N loading is significantly influenced the horizontal and vertical hetero- potentially going to transfer marine ecosystems from N lim- geneities of phytoplankton P deficiency in the Changjiang itation to P limitation. Seasonal P limitation has become ev- plume. Allochthonous APA was detected at nutrient-enriched ident in some estuarine and coastal ecosystems (Howarth freshwater end. Excessive N (∼ 10 to 112 µM) was observed and Marino, 2006). The altered nutrient dynamics will sub- throughout the entire plume surface. In the plume fringe fea- sequently influence ecosystem structure and function and turing stratification and excess N, diapycnal phosphate sup- threaten the safety of aquatic environments. Evidence also ply was blocked and phytoplankton APA was stimulated for showed that unbalanced nutrient composition might drive growth. We observed an upwelling just attaching to the tur- harmful algal growth (Varkitzi et al., 2010) and modulate car- bidity front at seaward side where Chl a peaked yet much bon (C) sequestration amount due to the adjustment of cellu- less APA was detected. An external phosphate supply from lar C / P ratios of plankton living in varying N / P conditions subsurface, which promoted phytoplankton growth but inhib- (Geider and Roche, 2002). The ambient nutrient concentra- ited APA, was suggested to be sourced from the Nearshore tions that determine the exchange rate between a cell and Kuroshio Branch Current. In the so hydrographically com- its medium may vary along the mixing gradient from river to plicated Changjiang plume, phosphate supply instead of its sea, as well as processes such as uptake, transformation, rem- concentration may be more important in determining the ex- ineralization, and diapycnal supply. Ecosystems’ response to pression of APA. Meanwhile, allochthonous APA may also nutrient loading has been demonstrated; however, the infor- alter the usefulness of APA as a P-stress indicator. mation about the dynamic nutrient pattern of excess N ver- sus the influence of nutrient stoichiometry on phytoplankton growth around riverine plumes and adjacent coastal ecosys- tems requires more exploration. 1 Introduction Under excess-N conditions, P generally limits phytoplank- ton growth in plumes and coastal areas where light is suf- Rapid urbanization in past decades along coastlines has intro- ficient. P limitation in ecosystems could be determined by duced massive loadings of nutrients and organics into rivers, Published by Copernicus Publications on behalf of the European Geosciences Union. 410 Y.-F. Tseng et al.: Phosphorus stress of coastal phytoplankton surrounding nutrient concentrations coupled with N / P ratios complicated CJ plume region. A conceptual model was con- (Krom et al., 1991; Dortch and Whitledge, 1992). However, structed and the properness of usage of various P-limitation most phytoplankton species have very high affinity with N indicators was discussed from different perspectives. and P; the ambient N or P concentration may have been be- low the analytical detection while growth is ongoing. Re- cently, a physiological indicator of phytoplankton P defi- 2 Materials and methods ciency – alkaline phosphatase (AP) activity (APA) – has been widely used (Mather et al., 2008; Duhamel et al., 2010). 2.1 Study area and sampling AP is an inducible extracellular enzyme generally existent Our cruise was conducted during 14–24 August of 2011, just in freshwater and marine plankton, which can cleave phos- about one week after the invasion of Super-Typhoon Muifa phate from dissolved organic phosphorus (DOP) molecules (27 July–9 August; the highest wind speed ∼ 175 km h−1; when surrounding or intracellular phosphate concentration see more details in Hsiao et al., 2013). A total of 28 sampling is low (Perry, 1972). Growing evidence has confirmed that stations were set in the field of 30–32◦ N and 121–124◦ E, phytoplankton can utilize DOP via generating AP under P where the CJ plume significantly influenced (Fig. 1). Five depletion (Hoppe, 2003). This characteristic makes APA a stations were deployed in the Qiantang (QT) estuary also. good indicator of P stress or P deficiency of phytoplankton Vertical profiles of temperature, salinity, and fluorescence in aquatic ecosystems (Dyhrman and Ruttenberg, 2006; Lo- were recorded with a CTD profiler (SBE-9/11 plus, SeaBird) mas et al., 2010). and packaged sensors at a sampling interval of 3 Hz. Water The East China Sea (ECS) is one of the largest marginal samples were collected starting from ∼ 3 m deep to the near- seas in the western North Pacific Ocean. According to his- bottom layer using a rosette sampler assembled with 10 L torical records, the ECS was considered as an N-limited Go-Flo bottles for chemical analyses. ecosystem as other marine ecosystems (Chen et al., 2001). The Changjiang (CJ) River holds large water discharge (an- 2.2 Nutrients, total suspended matter, and Chl a ∼ 3 −1 nual mean 924 km yr ) and high nutrient concentrations, analyses serving as the major nutrient source to the ECS (Zhang et al., 2007). Since the 1950s, rapidly growing anthropogenic ac- Nutrient water samples were filtered through Whatman tivities, such as fertilizer applications, have caused ∼ 8 × in- GF/F filters and analyzed immediately on board for DIN crease in N concentration and fluxes in the CJ River (Wang, − − + (DIN = NO3 + NO2 + NH4 ) and phosphate following the 2006). In addition, the CJ River was recorded to be a high standard colorimetric protocols (Parsons et al., 1984). The N / P ratio system (Liu et al., 2009) and “excess N” was found analyses were conducted with a continuous-flow injection in the surface water covering more than one-third of the ECS autoanalyzer (Technicon AA3-HR; SEAL Analytical). The in summer (Wong et al., 1998). Obviously, increased riverine detection limits for DIN and phosphate with 1 cm quartz nutrient loads have induced major changes of nutrient stoi- tubes were 0.1 µM and 0.05 µM, respectively. Chl a sam- chiometry in coastal zones (Chai et al., 2006), and it seems ples were filtered onto Whatman GF/F filters under pressure that the increasing trend and the expansion of influenced ar- < 100 mmHg and then kept in liquid N2 before analysis. In eas are non-stopping. Consequently, vast areas around the CJ the laboratory, Chl a samples were extracted in the dark at River mouth and ECS shelf are suggested to be P-limiting 4 ◦C with 100 % acetone for 12–16 h and then measured with for phytoplankton growth (Harrison et al., 1990; Wang et a fluorometer (10-AU; Turner Designs) following the method al., 2003). Likely, such P limitation will be more serious of Welschmeyer (1994). Total suspended matter (TSM) sam- in plume regions during the summer period, when river dis- ples were filtered onto Whatman GF/F filters, folded and charge and productivity are both promoted. wrapped by precombusted aluminum foils after salt washing, In this study, the Changjiang plume (CJ plume) was se- and then freeze-dried for 24 h for weighing in laboratory. lected to explore the consistency between chemical and phys- iological indicators, such as phosphate concentrations, N / P 2.3 Bulk APA ratios, and APA values, in representing a P-limitation envi- ronment. In addition, the Nearshore Kuroshio Branch Cur- Samples for bulk APA assays were filtered through a 100 µm rent (NKBC) brings nutrient-enriched subsurface water with mesh to remove large zooplankton and then collected with low N / P ratios up onto the ECS shelf even approaching the 2 L acid-cleaned polycarbonate bottles. The zooplankton- CJ estuary in summer (Yang et al., 2012, 2013). NKBC in- free samples were smoothly preconcentrated onto 0.2 µm trusion may compensate the excess N in the plume region polycarbonate filters and then resuspended in 50 mL filtered and further complicate the unbalanced nutrient status of the seawater (< 0.2 µm) from the same sampling depth. APA CJ plume system. In this study, we presented 3-D structures was measured using a sensitive fluorometric protocol similar of hydrographic parameters, nutrients, Chl a, and applied to Perry (1972) with 3-o-methylfluorescein phosphate sub- APA assays to probe whether phytoplankton physiologically strate (3-o-MFP; Sigma). For each sample, 6 mL triplicate endured P stress in such a N-replete and hydrographically sample waters and 0.75 mL 3-o-MFP substrate were mixed Biogeosciences, 11, 409–423, 2014 www.biogeosciences.net/11/409/2014/ Y.-F.