Alkaline phosphatase activity and its relationship to inorganic phosphorus in the transition zone of the North-western African upwelling system

The enzymatic activity of alkaline phosphatase (APA) was studied in the transition zone between the African upwelling system and the open ocean waters of the Canary Islands region. This region is recurrently dominated by the presence of upwelling filaments that may transport nutrient-enriched waters out into the open ocean before nutrients become exhausted by plankton. Turnover rates by APA were generally low in the whole region, but detectable in all the measurements carried out. On average, turnover rates were higher in the upwelling stations, and APA in those waters seemed to be mainly generated by heterotrophic bacteria to supply easily assimilable organic C. APA outside the upwelling area showed an inverse hyperbolic relationship with increasing phosphate, suggesting the presence of both constitutive and Pi-inducible APA. In these offshore waters, a threshold of 0.1 μM of phosphate could be defined for the regulatory function of Pi on APA. Thus, APA in nutrient-poor waters seemed to be induced to compensate for Pi-deficiency. Turnover rates in the filaments showed basal (probably constitutive) levels, whereas they increased in the surrounding waters, where phosphate concentration presumably did not satisfy plankton P-demands. The fertilising effect of the filaments and associated cyclonic eddies extended to at least 175 km offshore, where basal alkaline phosphatase activities were still found. The magnitude of this effect depends probably on the intensity of upwelling events and the degree of recirculation of filament water back to the coastal jet.     

Manganese and iron distributions off central California influenced by upwelling and shelf width

In July 2002, a combination of underway mapping and discrete profiles revealed significant along-shore variability in the concentrations of manganese and iron in the vicinity of Monterey Bay, California. Both metals had lower concentrations in surface waters south of Monterey Bay, where the shelf is about 2.5 km wide, than north of Monterey Bay, where the shelf is about 10 km wide. During non-upwelling conditions over the northern broad shelf, dissolvable iron concentrations measured underway in surface waters reached 3.5 nmol L⁻¹ and dissolved manganese reached 25 nmol L⁻¹. In contrast, during non-upwelling conditions over the southern narrow shelf, dissolvable iron concentrations in surface waters were less than 1 nmol L⁻¹ and dissolved manganese concentrations were less than 5 nmol L⁻¹. A pair of vertical profiles at 1000 m water depth collected during an upwelling event showed dissolved manganese concentrations of 10 decreasing to 2 nmol L⁻¹, and dissolvable iron concentrations of 12–20 nmol L⁻¹ in the upper 100 m in the north, compared to 3.5–2 nmol L⁻¹ Mn and 0.6 nmol L⁻¹ Fe in the upper 100 m in the south, suggesting the effect of shelf width influences the chemistry of waters beyond the shelf.These observations are consistent with current understanding of the mechanism of iron supply to coastal upwelling systems: Iron from shelf sediments, predominantly associated with particles greater than 20 μm, is brought to the surface during upwelling conditions. We hypothesize that manganese oxides are brought to the surface with upwelling and are then reduced to dissolved manganese, perhaps by photoreduction, following a lag after upwelling.Greater phytoplankton biomass, primary productivity, and nutrient drawdown were observed over the broad shelf, consistent with the greater supply of iron. Incubation experiments conducted 20 km offshore in both regions, during a period of wind relaxation, confirm the potential of these sites to become limited by iron. There was no additional growth response when copper, manganese or cobalt was added in addition to iron. The growth response of surface water incubated with bottom sediment (4 nmol L⁻¹ dissolvable Fe) was slightly greater than in control incubations, but less than in the presence of 4 nmol L⁻¹ dissolved iron. This may indicate that dissolvable iron is not as bioavailable as dissolved iron, although the influence of additional inhibitory elements in the sediment cannot be ruled out.     

Effects of variable winds on biological productivity on continental shelves in coastal upwelling systems

The production and distribution of biological material in wind-driven coastal upwelling systems are of global importance, yet they remain poorly understood. Production is frequently presumed to be proportional to upwelling rate, yet high winds can lead to advective losses from continental shelves, where many species at higher trophic levels reside. An idealized mixed-layer conveyor (MLC) model of biological production from constant upwelling winds demonstrated previously that the amount of new production available to shelf species increased with upwelling at low winds, but declined at high winds [Botsford, L.W., Lawrence, C.A., Dever, E.P., Hastings, A., Largier, J., 2003. Wind strength and biological productivity in upwelling systems: an idealized study. Fisheries Oceanography 12, 245–259]. Here we analyze the response of this model to time-varying winds for parameter values and observed winds from the Wind Events and Shelf Transport (WEST) study region. We compare this response to the conventional view that the results of upwelling are proportional to upwelled volume. Most new production per volume upwelled available to shelf species occurs following rapid increases in shelf transit time due to decreases in wind (i.e. relaxations). However, on synoptic, event time-scales shelf production is positively correlated with upwelling rate. This is primarily due to the effect of synchronous periods of low values in these time series, paradoxically due to wind relaxations. On inter-annual time-scales, computing model production from wind forcing from 20 previous years shows that these synchronous periods of low values have little effect on correlations between upwelling and production. Comparison of model production from 20 years of wind data over a range of shelf widths shows that upwelling rate will predict biological production well only in locations where cross-shelf transit times are greater than the time required for phytoplankton or zooplankton production. For stronger mean winds (narrower shelves), annual production falls below the peak of constant wind prediction [Botsford et al., 2003. Wind strength and biological productivity in upwelling systems: an idealized study. Fisheries Oceanography 12, 245–259], then as winds increase further (shelves become narrower) production does not decline as steeply as the constant wind prediction.     

Coupling of life cycles of the copepods Calanus chilensis and Centropages brachiatus to upwelling induced variability in the central-southern region of Chile

A time series of zooplankton sampling carried out at Station 18 off Concepción (36°S, 73°W) from August 2002 to December 2003 allowed the study of annual life cycles of the copepods Calanus chilensis and Centropages brachiatus in association with environmental variability in the coastal upwelling zone. Changes in the abundance of eggs, nauplii, and copepodids were assessed from samples taken at a mean time interval of ca. 20 days. Upwelling variability in near-surface waters was reflected in seasonal changes in salinity, water column stratification, and oxycline depth, as well as a weak seasonal signal in sea surface temperature (1-2 °C). Both copepods exhibited similar life cycles, characterized by continuous reproduction throughout the year. Estimates of generation times, as a function of temperature, were 25-30 days for C. chilensis and 27-35 days for C. brachiatus, predicting about 12 and 10 generations a year, respectively. These estimates were consistent with reproduction pulses observed in the field. It was thus suggested that copepods may grow under non-limiting food conditions in this upwelling area. However, despite continuous reproduction, there were abrupt changes in population sizes along with the disappearance of early naupliar and copepodid stages taking place even during the upwelling season (spring/summer). These changes were attributed to sudden increases in mortality taking place in spring or early summer, after which the populations remained at low levels through the fall and winter. It is thus suggested that, in addition to variability in the physical environment, biological interactions modulating changes in copepod mortality should be considered for understanding copepod life cycles in highly productive upwelling systems.     

Primary production in the eastern tropical Pacific: A review

The eastern tropical Pacific includes 28 million km² of ocean between 23.5°N and S and Central/South America and 140°W, and contains the eastern and equatorial branches of the north and South Pacific subtropical gyres plus two equatorial and two coastal countercurrents. Spatial patterns of primary production are in general determined by supply of macronutrients (nitrate, phosphate) from below the thermocline. Where the thermocline is shallow and intersects the lighted euphotic zone, biological production is enhanced. In the eastern tropical Pacific thermocline depth is controlled by three interrelated processes: a basin-scale east/west thermocline tilt, a basin-scale thermocline shoaling at the gyre margins, and local wind-driven upwelling. These processes regulate supply of nutrient-rich subsurface waters to the euphotic zone, and on their basis we have divided the eastern tropical Pacific into seven main regions. Primary production and its physical and chemical controls are described for each.Enhanced rates of macronutrient supply maintains levels of primary production in the eastern tropical Pacific above those of the oligotrophic subtropical gyres to the north and south. On the other hand lack of the micronutrient iron limits phytoplankton growth (and nitrogen fixation) over large portions of the open-ocean eastern tropical Pacific, depressing rates of primary production and resulting in the so-called high nitrate-low chlorophyll condition. Very high rates of primary production can occur in those coastal areas where both macronutrients and iron are supplied in abundance to surface waters. In these eutrophic coastal areas large phytoplankton cells dominate; conversely, in the open-ocean small cells are dominant. In a ‘shadow zone’ between the subtropical gyres with limited subsurface ventilation, enough production sinks and decays to produce anoxic and denitrified waters which spread beneath very large parts of the eastern tropical Pacific.Seasonal cycles are weak over much of the open-ocean eastern tropical Pacific, although several eutrophic coastal areas do exhibit substantial seasonality. The ENSO fluctuation, however, is an exceedingly important source of interannual variability in this region. El Niño in general results in a depressed thermocline and thus reduced rates of macronutrient supply and primary production. The multi-decadal PDO is likely also an important source of variability, with the ‘El Viejo’ phase of the PDO resulting in warmer and lower nutrient and productivity conditions similar to El Niño.On average the eastern tropical Pacific is moderately productive and, relative to Pacific and global means, its productivity and area are roughly equivalent. For example, it occupies about 18% of the Pacific Ocean by area and accounts for 22–23% of its productivity. Similarly, it occupies about 9% of the global ocean and accounts for 10% of its productivity. While representative, these average values obscure very substantial spatial and temporal variability that characterizes the dynamics of this tropical ocean.     

The circulation of the eastern tropical Pacific: A review

During the 1950s and 1960s, an extensive field study and interpretive effort was made by researchers, primarily at the Scripps Institution of Oceanography, to sample and understand the physical oceanography of the eastern tropical Pacific. That work was inspired by the valuable fisheries of the region, the recent discovery of the equatorial undercurrent, and the growing realization of the importance of the El Niño phenomenon. Here we review what was learned in that effort, and integrate those findings with work published since then as well as additional diagnoses based on modern data sets.Unlike the central Pacific, where the winds are nearly zonal and the ocean properties and circulation are nearly independent of longitude, the eastern tropical Pacific is distinguished by wind forcing that is strongly influenced by the topography of the American continent. Its circulation is characterized by short zonal scales, permanent eddies and significant off-equatorial upwelling. Notably, the Costa Rica Dome and a thermocline bowl to its northwest are due to winds blowing through gaps in the Central American cordillera, which imprint their signatures on the ocean through linear Sverdrup dynamics. Strong annual modulation of the gap winds and the meridional oscillation of the Intertropical Convergence Zone generates a Rossby wave, superimposed on the direct forcing, that results in a southwestward-propagating annual thermocline signal accounting for major features of observed thermocline depth variations, including that of the Costa Rica Dome, the Tehuantepec bowl, and the ridge–trough system of the North Equatorial Countercurrent (NECC). Interannual variability of sea surface temperature (SST) and altimetric sea surface height signals suggests that the strengthening of the NECC observed in the central Pacific during El Niño events continues all the way to the coast, warming SST (by zonal advection) in a wider meridional band than the equatorially trapped thermocline anomalies, and pumping equatorial water poleward along the coast.The South Equatorial Current originates as a combination of equatorial upwelling, mixing and advection from the NECC, and Peru coastal upwelling, but its sources and their variability remain unresolved. Similarly, while much of the Equatorial Undercurrent flows southeast into the Peru Undercurrent and supplies the coastal upwelling, a quantitative assessment is lacking. We are still unable to put together the eastern interconnections among the long zonal currents of the central Pacific.     

The influence of the coast on the dynamics of upwelling fronts: Part II. Numerical simulations

We investigated the dynamics of upwelling fronts near a coast. This work was first motivated by laboratory experiments [Bouruet-Aubertot, Linden, Dyn. Atmos. Oceans, 2002] in which the front is produced by the adjustment of a buoyant fluid initially confined within a bottomless cylinder. It was shown that cyclonic eddies consisting of coastal waters are enhanced when the front is unstable near the coast (the outer vertical boundary). The purpose of this paper is to provide further insights into this process. We reproduced the experimental configuration using a three-dimensional model of the primitive equations. We first show that for coastal fronts more potential energy, in terms of the maximum available potential energy, is released than for open-ocean fronts. Therefore, waves of larger amplitude are generated during the adjustment and the mean flow that establishes has a higher kinetic energy in the former case. Then as baroclinic instability starts and wave crests reach the boundary, cyclonic eddies are enhanced as in the laboratory experiments and in a similar way. However, in contrast to the laboratory experiments, offshore advection of cyclonic eddies can occur in two stages, depending on the spatial organization of the baroclinic wave. When the baroclinic wave consists of the sum of different modes and is thus highly asymmetric, the offshore advection of cyclonic eddies occurs just after their enhancement at the boundary, as in the laboratory experiments. By contrast, when a single-mode baroclinic wave develops, neighboring cyclonic eddies first merge before being advected offshore. Very different behavior is observed for open-ocean fronts. First a mixed baroclinic–barotropic instability grows. Then the eddies transfer their energy to the mean flow and the barotropic and baroclinic instabilities start again. An excellent agreement is obtained with the main result obtained in the laboratory experiments: the ratio between growth rates of surface cyclonic and anticyclonic vorticity increases as the instability develops nearer to the coast.     

Possible refugia for reefs in times of environmental stress

This paper investigates the refuge potential of (1) upwelling areas, (2) coral areas at medium depth, and (3) offshore bank and island reefs in a scenario of increased global warming, and thus increased sea surface temperature (SST) and increased solar UV radiation. (1) Observations on coral health and water temperature in the subtropical Atlantic (Eleuthera and Cat Island, Bahamas) and Indian Ocean (Sodwana Bay, South Africa) suggest a link between cool water delivered by upwelling and coral health. After the 1998 bleaching event, caused by strong SST anomalies, coral health and recovery from the previous year's bleaching was significantly better on the narrow southern Cat Island shelf (70% of corals healthy) where the presence of cold water was observed, which was attributed to small-scale upwelling, than on the wide northern Eleuthera shelf (44% of corals healthy), where downwelling of hot bank waters was believed to have damaged corals. In South Africa, regular, short-term upwelling events in five summers reduced SST to well below bleaching level. (2) In the northern Red Sea (Safaga Bay) and in South Africa (Sodwana Bay), wide areas with either coral frameworks or non-framework communities exist. Calculations show that if the top 10 m (20 m) of the ocean became inhospitable to corals, still 50.4% (17.5%) of the coral area would remain intact in the Red Sea and 99% (40%) in South Africa. (3) Offshore bank and island reefs investigated in the Turks, Caicos, and Mouchoir Banks and Grand and Little Cayman showed high rates of mortality and coral diseases. The most remote sites (Mouchoir Bank) were not the healthiest. Refuge areas appear to exist in (1) and (2), but in (3) only if vigorous water-circulation is encountered.     

沿岸上升流和沿岸急流的一个半解析理论

在考虑了陆架地形后，在垂直海岸的x z剖面上 ，对Boussinesq流体的非线性海洋运动方程求得了总动量守恒、温度守恒和位势涡度守恒的 普 适形式，进而得到流函数所满足的椭圆型二阶偏微分方程，在给定流体沿地形运动的条件下 ，算出问题的解. 计算结果表明，沿岸可以出现上升流也可以出现下沉流，它依赖于海洋的 大尺度背景条件. 计算所得的上升流、沿岸急流、温度的锋区结构与一些观测事实接近.     

极隙区电子束流模电磁不稳定性

解析地研究了中高度（离他心3－4个地球半径）极隙区极低密度上行电子束流引起的沿磁力线传播的电磁个稳定性，上行电子束流和背景等离子体都考虑成冷等离子体．结果表明，上行电子引起的左旋和右旋圆偏振电磁来流模是不稳定的，当它与离子回旋模耦合时增长率达到最大值，频率色散关系仍为电子束流模特征这此结果对解释权隙区纬度地面站低频电磁波观测资料和理解极隙区动力学过程是很有益的．