长江口上升流区营养盐的分布及其通量的初步估算

根据2004年5月份长江口调查资料,分析了春季长江口上升流现象及其对营养盐分布的影响;初步估算了春季上升流的营养盐通量。结果表明:在春季,低温、高盐、低溶解氧的上升流稳定存在于122°20′～123°00′E,31°00′～32°00′N海区的10m 层和底层之间,并可以涌升到10m 层以上海区。上升流为上层海区输入了丰富的 PO_4-P 和相对低浓度的 NO_3-N 和 SiO_3-si。对上升流营养盐通量的计算表明,春季上升流中磷酸盐输送通量远高于长江径流输入,可能会成为影响该海区磷酸盐分布以及浮游植物生长的一个值得关注的因素:而氮和硅营养盐则不如长江径流输入量大。     

Possible mixotrophy of pigmented nanoflagellates: Microbial plankton biomass, primary production and phytoplankton growth in the NW Iberian upwelling in spring

Microbial plankton biomass, primary production (PP) and phytoplankton growth rates (μ) were estimated along the NW Iberian margin during an upwelling relaxation event. Although the interaction between wind forcing and coastline singularities caused high spatial variability in PP (0.4-8.4 g C m⁻² d⁻¹), two domains (coastal and oceanic) could be distinguished regarding microbial plankton biomass and μ. At the coastal domain, with higher influence of upwelling, diatoms showed an important contribution (27 ± 17%) to total autotrophic biomass (AB). Nonetheless, AB was dominated by autotrophic nanoflagellates (ANF) at both realms, accounting for 62 ± 16% and 89 ± 6% of the integrated AB at the coastal and oceanic domain respectively. AB and heterotrophic biomass (HB) were significantly higher at the oceanic than at the coastal domain, with both biomasses covarying according to HB:AB = 0.33. Whereas the low phytoplankton carbon to chlorophyll a ratio (C_ph:chl a = 38 ± 3) and the high μ = 0.54 ± 0.09 d⁻¹ registered at the coastal stations suggest that phytoplankton was not nutrient limited at this domain, the values (C_ph:chl a = 157 ± 8; μ = 0.17 ± 0.02 d⁻¹) recorded at the oceanic domain point to severe nutrient limitation. However, the high Fv/Fm fluorescence ratios (0.56 ± 0.09) measured at the sea surface in the oceanic domain suggest that nutrient limitation did not occur. To reconcile these two apparently opposite views, it is suggested the occurrence of mixotrophic nutrition of ANF, with heterotrophic nutrition supplying about 75% of carbon requirements.     

Off-shelf fluxes of labile materials by an upwelling filament in the NW Iberian Upwelling System

Daily changes in the concentrations of carbon and nitrogen species were monitored during the course of a Lagrangian drifter experiment in a recurrent upwelling filament south of Cape Finisterre (NW Iberian Upwelling System). A drifting buoy released at the southern edge of the upwelling centre generated by the Cape moved 60km southwestwards from 3 to 7 August 1998. Organic matter in the 50m deep study water mass (average 77±2 μM C) consisted of: 57μM C of dissolved organic matter (DOM) with a C/N molar ratio of 19±2; 6μM C of DOM with a C/N ratio of 9±2, and 14μM C of 50% DOM and 50% suspended organic matter (POM_susp) with a C/N ratio of 6.0±0.4.Net conversion of consumed inorganic salts into accumulated TOM=POM_susp+DOM was 40% for nitrogen and 30% for carbon. Since the parcel of water crossed the shelf-edge, these conversion efficiencies are equivalent to net horizontal export-ratio of 0.4 and 0.3 respectively. A second drifter was deployed in the offshore-end of the filament, and was displaced 20km west between 14 and 17 August 1998. Nitrate was exhausted in the surface water and no significant changes were observed in the variables measured during the course of the second experiment. Low C/N ratios (6.5±0.4) and rapid POM_susp/DOM inter-conversion in the 20 μM C excess observed in the study volume points to the persistence of the labile materials formed on the shelf during transport to the ocean. Our data demonstrate

a) the key role of upwelling filaments in off-shelf export of organic materials and

b) the major contribution of DOM to this horizontal export, a previously unaccounted amount.

The high nitrogen content of the materials exported make them attractive organic substrates for use by microbial populations in the adjacent oligotrophic ocean.     

Simulation studies of trophic flows and nutrient cycles in Benguela upwelling foodwebs

The traditional model of short, efficient food chains in upwelling regions is revised to incorporate microplanktonic processes and near-exclusive carnivory by anchovy. The new model is based on analyses of outputs of simulation models. These include a size-based micro/mesoplankton model of a Benguela upwelling community and a model of euphausiid grazing. On average, large-celled phytoplankton are responsible for only one-quarter to one-third of total primary production, the remainder occurring by cells <25 μm. Trophic transfers of carbon in the models take place chiefly via one or two steps, after which carbon is lost through respiration and the sinking of faecal material. Trophic efficiencies can be high, generally 40–50 per cent for the first transfer, 15–20 per cent for the second, and 5–10 per cent for remaining transfers. A first attempt is made at linking two models of processes occurring on different scales of time and space. Of the problems encountered, potentially the most difficult to overcome appears to be the manner in which feedbacks can be incorporated from the large to the small scale. C:N ratios of predator and prey are important in determining C:N ratios of faecal material. In a model of a euphausiid population, these ratios vary from 6 to 17 depending on whether animal or plant material respectively is being consumed. This is important in determining rates of carbon export from surface waters through sinking of faecal pellets. The effect of the mesoscale physical environment on plankton distributions and transport needs to be addressed through modelling studies, in order to link planktonic processes to macrozooplankton- and fish-feeding.     

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.     

Hydrographic and atmospheric analysis of an autumnal upwelling event in the Ria of Vigo (NW Iberian Peninsula)

An autumnal upwelling event was observed in the Ria of Vigo (NW Iberian Peninsula) on 15th November 2001. This event was analyzed by means of thermohaline variables measured at CTD stations located in the study area, satellite sea surface temperature and wind data provided by QuikSCAT. Salinity and temperature distributions revealed that the upwelled water mass was Eastern North Atlantic Central Water (ENACW), typically observed during summer upwelling events. However, previous to the upwelling event, the characteristic autumnal body of water was recorded on 31st October 2001. Subsequent sea surface temperature and upwelling index corroborated the presence of an autumnal upwelling event of approximately 15 days duration. The probability of upwelling-favorable winds (from 1999 to 2004) was lower during autumn–winter than during spring–summer, although they may occur at any time of the year. Probabilities of 45% were calculated for February and November, with the highest probability (65%) corresponding to July.     

Physical drivers of phytoplankton production in the southern Benguela upwelling system

Investigations of primary production (PP) were undertaken in the southern Benguela ecosystem during two research surveys in October 2006 and May 2007. Significant differences in environmental conditions, as well as biomass and PP, were observed between October and May. During October, integrated biomass and PP were significantly higher, ranging from 20.43 to 355.01 mg m⁻², and 0.71 to 6.98 g C m⁻² d⁻¹, respectively, than in May, where the range was 47.92–141.79 mg m⁻², and 0.70–3.35 g C m⁻² d⁻¹, respectively. Distribution patterns indicated low biomass and PP in newly upwelled water along the coast, higher biomass and PP in the mid-shelf region, while lower values were observed at and beyond the shelf edge. Latitudinal variations showed consistently higher biomass and PP in the St. Helena Bay region compared to biomass and PP south of Cape Town. During both surveys, phytoplankton communities were comprised primarily of diatoms and small flagellates, with no significant differences. Phytoplankton adaptation to environmental variability was characterised by increased P_m^B and E_k under elevated temperatures and irradiance, while no clear relationships were evident for α^B. Generalised Additive Models (GAMs) showed that photosynthetic parameters were all significant predictors of photosynthesis rates (P_z), with P_m^B being the most important, accounting for 36.97% of the deviance in P_z. However, biomass levels and environmental conditions exerted a much greater influence on P_z, with irradiance explaining the largest proportion (68.24%) of the deviance. Multiple predictor GAMs revealed that 96.26% of the deviance in P_z could be explained by a model which included nitrate, chlorophyll a, and irradiance.     

Responses of nutrient biogeochemistry and nitrogen cycle to seasonal upwelling in coastal waters of the eastern Hainan Island

The coastal upwelling has profound influence on the surrounding ecosystem by supplying the nutrient-replete water to the euphotic zone. Nutrient biogeochemistry was investigated in coastal waters of the eastern Hainan Island in summer 2015 and autumn 2016. From perspectives of nutrient dynamics and physical transport, the nutrient fluxes entered the upper 50 m water depth（between the mixed layer and the euphotic zone） arisen from the upwelling were estimated to be 2.5-5.4 mmol/（m²·d）,...     

Microzooplankton feeding impact in a coastal upwelling system on the NW Iberian margin: The Ría de Vigo

The dilution technique, combined with identification and enumeration of pico-, nano- and micro-plankton by microscopy, was used to estimate microzooplankton impact on the microbial community in surface waters of a coastal embayment on the NW Iberian upwelling system. Microzooplankton were important consumers of autotrophic and heterotrophic plankton in this system, feeding up to 93% of standing stock and more than 100% of production of several groups. Heterotrophic bacteria and heterotrophic picoflagellates experienced the highest and constant impact, with 75–84% of their standing stocks and 85–102% of their production being channelled through the microbial food web. Pico- and nano-phytoplankton were also consumed, although maximum grazing occurred on diatoms during upwelling events, coinciding with highest primary production. Predation on pico-nano-heterotrophs was especially relevant under downwelling conditions, when consumption of total carbon and particularly autotrophic carbon was considerably lower than during upwelling. The results suggest that the existence of a multivorous food web, extending from the microbial loop to the herbivorous food web, could be a major feature in this coastal upwelling system. The microbial loop, which occurs as a permanent background in the system, would contribute to sustain the microbial food web during downwelling, whereas the herbivorous food web could coexist with a microbial food web based on large diatoms during upwelling. The multivorous food web would partially divert diatoms from sinking and hence favour the retention of organic matter in the water column. This could enhance the energy transfer to higher pelagic trophic levels in coastal upwelling systems.     

Consequences of winter upwelling events on biogeochemical and phytoplankton patterns in a western Galician ria (NW Iberian peninsula)

The consequences of two upwelling events in mid- (MW) and late (LW) winter on biogeochemical and phytoplankton patterns were studied in the Pontevedra Ria and compared with the patterns measured under typical winter conditions and under a summer upwelling event. Thermohaline patterns measured during the mid-winter upwelling event (MW-up) revealed the intrusion of saltier seawater (35.9) into the ria associated with the Iberian Poleward Current (IPC). During the late-winter upwelling event (LW-up), the seawater which had welled up into the ria showed characteristics of the Eastern North Atlantic Central Water mass (ENACW). In both cases the measured water residence time (4 days during MW-up and 10 days during LW-up) was related to both meteorological and fluvial forcing. This residence time contrasts with that of summer upwelling (7 days) and with that estimated under unfavorable upwelling atmospheric conditions (2–4 weeks). During MW-up, the ria became poor in nutrients due to continental freshwater dilution, associated with the shorter residence time of the water, and the intrusion of IPC, which is a water body poor in nutrient salts: 2.9 μM of nitrate, 0.1 μM of phosphate and 1.5 μM of silicate. During this event, the ria exported 3.4 mol_DIN s⁻¹, compared with 6.9 mol_DIN s⁻¹ in non-upwelling conditions. Phytoplankton showed a uniform distribution throughout the ria, as during unfavorable upwelling conditions, and was characterized by the dominance of diatoms, mainly Nitzschia longissima and Skeletonema costatum. During LW-up, a nutrient depletion in the photic layer also occurred, but as a result of a phytoplankton spring bloom developing at this time. The ria was a nutrient trap where 4.1 mol_DIN s⁻¹ were processed by photosynthesis. This budget is three times higher than the one under non-upwelling conditions. In contrast with the MW-up, which had no effect on primary production, during LW-up the ria became more productive, although not as productive as during a summer upwelling event (9.9 mol_DIN s⁻¹). The taxonomic composition of the phytoplankton community did not change noticeably during LW-up and the summer upwelling, with the same species present and changing only in relative proportions. Diatoms were always the dominant microphytoplankton community, with Pseudonitzschia pungens, Thalassionema nitzschioides and several species of Chaetoceros as characteristic taxons.     

On the reconstruction of upwelling history: Namibia upwelling in context

Eastern boundary upwelling is an important aspect of the modern ocean, despite the relatively small area involved. Consequences of increased upwelling during the Neogene, as a result of intensifying trade winds, include increased heat transport by the ocean, increased mixing, increased fractionation of phosphate from silicate, and opportunities for evolution of new species. Coastal upwelling has been studied since the first decade of the 20th century. The processes involved are complex, and differ depending on geographic setting. Off Namibia, upwelling history has been studied by a number of drilling expeditions; the last of these was ODP Leg 175, which occupied seven sites on the continental slope between Walvis Ridge and Cape Town, during September and October of 1997. Productivity proxies all along southwestern Africa suggest the presence of increased upwelling and organic matter supply to the sea floor during glacial periods. At the same time, there is a decrease in the supply of diatoms and other siliceous plankton remains. This is the Walvis Opal Paradox, established through the contrasting results of studies by Diester-Haass (1985) and Oberhänsli (1991). We propose that the Walvis Opal Paradox is fundamental to the understanding of glacial–interglacial productivity fluctuations on a global scale. Furthermore, a central feature of the history of late Neogene upwelling off Namibia is the Matuyama Opal Maximum, centered between Gauss and Olduvai magnetic chrons [Wefer et al., Proc. ODP 175 (1998)]. It is due to the fact that diatom supply first increases (during the Gauss) when the planet cools and then decreases again during additional cooling, on entering the Quaternary. On a 400 000-year scale, peak productions are coincident with (or slightly lag) maximum seasonal contrast potential in the high-latitude insolation curve. We suggest that this is further evidence that the nutrient content of thermocline waters was diminished during glacial periods. The reasons why this should be so remain to be discovered.     

南海主要上升流及其与渔场的关系

<正>南海是我国的重要渔业产区,目前我国每年在这一海区的捕捞产量大约3×106 t。无论在南海北部还是在南海中部和南部海域都分布有优良渔场,这些渔场往往与上升流存在着密切关系。上升流是一种海水垂直向上的运动现象,通常因表层水体辐散所致,是海洋环流中的重要组成部分。上升流涌升速度与水平流速相比甚小,一般只有10–4~10–2 cm/s[1]。上升流可以把底层营养盐带到表层,为浮游植物的生长提供物质基础,进而为浮游动物、鱼类、虾类等     

Wintertime rates of net primary production and nitrate and ammonium uptake in the southern Benguela upwelling system

The elevated levels of primary productivity associated with eastern boundary currents are driven by nutrient- rich waters upwelled from depth, such that these regions are typically characterised by high rates of nitrate-fuelled phytoplankton growth. Production studies from the southern Benguela upwelling system (SBUS) tend to be biased towards the summer upwelling season, yet winter data are required to compute annual budgets and understand seasonal variability. Net primary production (NPP) and nitrate and ammonium uptake were measured concurrently at six stations in the SBUS in early winter. While euphotic zone NPP was highest at the stations nearest to the coast and declined with distance from the shore, a greater proportion was potentially exportable from open-ocean surface waters, as indicated by the higher specific nitrate uptake rates and f-ratios (ratio of nitrate uptake to total nitrogen consumption) at the stations located off the continental shelf. Near the coast, phytoplankton growth was predominantly supported by ammonium despite the high ambient nitrate concentrations. Along with ammonium concentrations as high as 3.6 µmol l^–1, this strongly suggests that nitrate uptake in the inshore SBUS, and by extension carbon drawdown, is inhibited by ammonium, at least in winter, although this has also been hypothesised for the summer.     

Nutrient controls on new production in the Bodega Bay, California, coastal upwelling plume

A theoretical framework for the time-dependent processes leading to the high rates of new production in eastern boundary upwelling systems has been assembled from a series of past upwelling studies. As part of the CoOP WEST (Wind Events and Shelf Transport) study, new production in the Bodega Bay upwelling area and it's control by ambient nitrate and ammonium concentrations and the advective wind regime are described. Data and analyses are focused primarily on the WEST 2001 cruise (May–June 2001) when the two legs differed greatly in wind regimes but not nutrient concentrations. Elevated concentrations of ammonium in upwelled water with high nitrate were observed in both legs. Nitrate uptake by phytoplankton as a function of nitrate concentration was linear rather than Michaelis–Menten-like, modulated by inhibitory levels of ammonium, yielding coefficients that enable the specific nitrate uptake element of new production to be estimated from nutrient concentrations. The range of specific nitrate uptake rates for the two legs of WEST 2001 were similar, essentially a physiological response to nutrient conditions. However, the low “realization” of new production i.e. incorporation of biomass as particulate nitrogen that occurred in this system compared to the theoretical maximum possible was determined by the strong advective and turbulent conditions that dominated the second leg of the WEST 2001 study. These data are compared with other upwelling areas using a physiological shift-up model [Dugdale, R.C., Wilkerson, F.P., Morel, A. 1990. Realization of new production in coastal upwelling areas: a means to compare relative performance. Limnology and Oceanography 35, 822–829].     

Degree of nutrient consumption as an aging index of upwelling or vertically mixed water in the northern Taiwan Strait

In order to determine the relative aging slams of upwelled or vertically mixed water in the northern Taiwan Strait, a recently developed method of calculating the degree of nutrient consumption (DNC) was employed. Upwelling was detected at a water depth of about 75m in summer. Bottom waters in the aphotic zone and newly-upwelled waters in the euphotic zone were found to be low in terms of their DNC. In general, a low DNC was noted alongside the other traditional upwelling indicators, such as lower temperature and degree of oxygen saturation, but higher salinity, apparent oxygen utilization, nutrient contents and chlorophytt-a. Enhanced vertical mixing, but without an apparent upwelling signal, was detected near the same location in winter.     

A comparison between the megafauna communities on the N.W. Iberian and Celtic continental margins—effects of coastal upwelling?

Megafauna biomass and feeding guilds were studied on the NW Iberian upwelling Continental Margin in order to determine the presence of enriched zones pointing to enhanced particle input. We compare these findings with similar data obtained from a transect across the Celtic Continental Margin that represents a regime without coastal upwelling. Additionally sediment concentrations of phytopigments (chlorophyll-a, phaeophorbides) representing recent inputs of algal production and of nucleic acids (DNA, RNA) are used as proxies for microbial biomass, to assess if there was a relation between these parameters and the megafauna distribution. The sediment on the upper slope (<1600 m) of the Iberian Margin was found to be inhabited by filter-feeding megafauna (26–73% of total invertebrate density, and 1–35% of biomass), and contained relatively low levels of phytopigments (3–6 ng/cm³ chlorophyll-a) and nucleic acids (12–16 μg⁻¹ DNA, 1.5–3.5 μg⁻¹ RNA). In contrast, on the upper slope of the Celtic Margin the dominant component of the megafauna were deposit-feeders (57–92% of total invertebrate density, and 23–90% of biomass) and the sediments contained higher concentrations of phytopigments and nucleic acid. These observations, supplemented by video records revealing the presence of current ripples on the Iberian upper slope, show that these upper slope regions are non-depositional, high energy environments. Conditions at the lower slope and the abyssal plain on the Iberian transect were more quiescent with large deposit-feeding holothurians dominating the megafauna (72–94% of invertebrate biomass), and with relatively high sediment concentrations of phytopigments (7–9 ng/cm³ chlorophyll-a, 157–170 ng/cm³ phaeophorbides) and nucleic acids (21–38 μg⁻¹ DNA, 2.4–5.5 μg⁻¹ RNA). On the basis of our data we argue that the benthic food for the deepest stations on the Iberian transect does not consist of shelf derived organic matter. More likely, fast sinking offshore blooms, possibly associated with filaments of upwelling water, form the major contribution to the annual food supply of the deep living megafauna.     

Seasonal and sub-seasonal oxygen and nutrient fluctuations in an embayment of an eastern boundary upwelling system: St Helena Bay

Seasonal, sub-seasonal and spatial fluctuations in bottom dissolved oxygen (DO) were examined in St Helena Bay, South Africa’s largest and most productive embayment, between November 2013 and November 2014. Alongshore bay characteristics were assessed through comparison of variables along the 50-m depth contour. A mean coefficient of variation of 0.35 provided a measure of the relative variability of near-bottom DO concentrations along this contour. Consistently lower DO concentrations in the southern region of the bay in summer and autumn are attributed to enhanced retention. Across-shelf transects captured the seasonal development of hypoxia in relation to the distribution of phytoplankton biomass. Exceptional dinoflagellate blooms form extensive subsurface thin layers preceding the autumn DO minima in the south of the bay, prior to winter ventilation of the bottom waters. The seasonal decline in DO concentrations in the bottom waters was marked by sub-seasonal events of hypoxia, and ultimately anoxia linked to episodic deposition of organic matter, as indicated by increases in bottom chlorophyll-a concentrations. Seasonal changes in bottom water macronutrient concentrations followed trends in apparent oxygen utilisation (AOU), both of which mirrored DO concentrations. In the south of the bay, nitrogen loss through denitrification/anammox in suboxic waters was indicated by a dissolved inorganic N deficit in the bottom waters, which was most pronounced in autumn.     

Weak and strong constraint data assimilation in the inverse Regional Ocean Modeling System (ROMS): Development and application for a baroclinic coastal upwelling system

We describe the development and preliminary application of the inverse Regional Ocean Modeling System (ROMS), a four dimensional variational (4DVAR) data assimilation system for high-resolution basin-wide and coastal oceanic flows. Inverse ROMS makes use of the recently developed perturbation tangent linear (TL), representer tangent linear (RP) and adjoint (AD) models to implement an indirect representer-based generalized inverse modeling system. This modeling framework is modular. The TL, RP and AD models are used as stand-alone sub-models within the Inverse Ocean Modeling (IOM) system described in [Chua, B.S., Bennett, A.F., 2001. An inverse ocean modeling system. Ocean Modell. 3, 137–165.]. The system allows the assimilation of a wide range of observation types and uses an iterative algorithm to solve nonlinear assimilation problems. The assimilation is performed either under the perfect model assumption (strong constraint) or by also allowing for errors in the model dynamics (weak constraints). For the weak constraint case the TL and RP models are modified to include additional forcing terms on the right hand side of the model equations. These terms are needed to account for errors in the model dynamics.Inverse ROMS is tested in a realistic 3D baroclinic upwelling system with complex bottom topography, characterized by strong mesoscale eddy variability. We assimilate synthetic data for upper ocean (0–450 m) temperatures and currents over a period of 10 days using both a high resolution and a spatially and temporally aliased sampling array. During the assimilation period the flow field undergoes substantial changes from the initial state. This allows the inverse solution to extract the dynamically active information from the synthetic observations and improve the trajectory of the model state beyond the assimilation window. Both the strong and weak constraint assimilation experiments show forecast skill greater than persistence and climatology during the 10–20 days after the last observation is assimilated.Further investigation in the functional form of the model error covariance and in the use of the representer tangent linear model may lead to improvement in the forecast skill.     

Modelling of upwelling and downwelling in the ocean

Numerical experiments on the reconstruction of upwelling and downwelling at the eastern boundary of the ocean were carried out in the framework of a multilayer model of the ocean involving the upper mixed layer (UML). The peculiarities of these phenomena when they are formed and attenuated owing to the strong intensification and abatement of the longshore wind have been studied. It is shown that cold waters are always involved from the thermocline to the UML during upwelling. In downwelling, this occurs as a rule. However, during upwelling the abatement of the wind may result in subduction —the inflow of warm waters from the UML to the thermocline.Translated by Mikhail M. Trufanov.