Chlorophyll <Emphasis Type="Italic">a</Emphasis> as a measure of seasonal coupling between phytoplankton and the monsoon periods in the Gulf of Oman

We present data from a long time-series study to describe the factors that control phytoplankton population densities and biomass in the coastal waters of Oman. Surface temperature, salinity, nutrients, dissolved oxygen, chlorophyll a (Chl a), and phytoplankton and zooplankton abundance of sea water were measured as far as possible from February 2004 through February 2006, at two stations along the southern coast of the Gulf of Oman. The highest concentrations of Chl a (3 mg m⁻³) were recorded during the southwest monsoon (SWM) when upwelling is active along the coast of Oman. However, results from our study reveal that the timing and the amplitude of the seasonal peak of Chl a exhibited interannual variability, which might be attributed to interannual differences in the seasonal cycles of nutrients caused either by coastal upwelling or by cyclonic eddy activity. Monthly variability of SST and concentrations of dissolved nitrate, nitrite, phosphate, and silicate together explained about 90% of the seasonal changes of Chl a in the coastal ecosystem of the Gulf of Oman. Phytoplankton communities of the coastal waters of Oman were dominated by diatoms for most part of the year, but for a short period in summer, dinoflagellates were dominant.     

Decadal weakening of the wind-induced upwelling reduces the impact of nutrient pollution in the Bay of Málaga (western Mediterranean Sea)

The hypothesis that nitrate versus phosphate regulates the coastal primary production has been assessed at different time scales in the northwest Alboran Sea. Time series of temperature, salinity, nutrients and chlorophyll a obtained at three stations located off Málaga city (the greatest coastal urban core along the Alboran Sea shoreline) from 1992 to 2006 have been analysed. At the decadal scale, temperature increased linearly while salinity decreased. These changes were related to a shift in the wind regime suggesting that coastal upwelling became steadily weaker. In contrast to phosphate, nitrate was positively correlated with salinity at the seasonal scale and decreased linearly from 1992 to 2006. Seasonal and decadal changes in chlorophyll a were correlated with nitrate (and uncorrelated with phosphate). However, non-regular variability in chlorophyll a was correlated with phosphate. Consequently, the results demonstrate that nitrate controls the phytoplankton biomass at the inter-annual scale while both nitrate and phosphate do so at a shorter time scale. The Bay of Málaga receives elevated entries of domestic waste waters that release high loads of phosphate compared to nitrate. Our analysis indicates that the expected impact of this pollution on chlorophyll a at the inter-annual scale is reduced in comparison with the effects of atmospheric forcing.     

Effects of Mississippi River water on phytoplankton growth and composition in the upper Barataria estuary,Louisiana

Diversion of river waters to adjacent estuaries may occur during wetland restoration, navigation channel development, or storms. We proposed that diversions of nitrogen- and phosphorus-enriched waters from the river to estuarine waters would result in increased phytoplankton biomass and shifts to noxious or harmful algal blooms. We tested this hypothesis by conducting four seasonal microcosm experiments in which Mississippi River water was mixed at different volume ratios with ambient estuarine waters of three lakes in the upper Barataria Basin, Louisiana, USA. These lakes included two brackish lakes that were in the path of diverted Mississippi River water, and a freshwater lake that was not. The results from the 3- to 8-day experiments yielded a predictable increase in phytoplankton biomass related to nutrient additions from Mississippi River water. The subsequent decreases in the dissolved nitrate + nitrite, soluble reactive phosphorus, and silicate concentrations explained 76 to 86% of the increase in chlorophyll a concentrations in the microcosms. Our experiments showed that cyanobacteria can successfully compete with diatoms for N and P resources even under non-limiting Si conditions and that toxic cyanobacteria densities can increase to bloom levels with increased Mississippi River water inputs to ambient waters in the microcosms. Diversions of Mississippi River into adjacent estuarine waters should be considered in relation to expected and, possibly, unexpected changes in phytoplankton communities to the receiving waters and coastal ecosystems.

     

GIS支持下考洲洋养殖水域浮游植物数量的时空分布及其与营养盐的相关性研究

在GIS和统计分析软件SPSS的支持下,利用数据插值、相关性分析和回归分析等方法,研究了枯水期和丰水期考洲洋养殖水域浮游植物数量的分布趋势及其与海水营养盐的相关关系。结果表明,枯水期浮游植物数量的分布较为均匀,其密集区分布于吉隆河口及其附近水域;而丰水期的数量变化幅度较大,密集分布区出现在湾口至盐洲岛北部水域,呈现出从湾西北部至湾口水域逐渐增加的分布趋势。两次调查相比,丰水期浮游植物的数量明显高于枯水期。相关性研究结果表明,调查期间浮游植物数量与无机氮和无机磷的相关性均不显著,但与活性硅酸盐呈显著相关,在枯水期两者之间呈显著正相关,而在丰水期则刚好相反,两者之间呈显著负相关。     

Nearshore abundance of dissolved and sedimented forms of phosphorus

Different forms of dissolved and sediment phosphates, both adsorbed and interstitial, were determined in the highly productive coastal waters of the Indian Ocean. Both inorganic and organic phosphate of the above fractions covered in this paper reveal the extent of inter-dependency as well as exemplify the role of sediment based nutrients to influence the biorhythm in the overlying waters. The area of study in the mudbank regions of Kerala coast, (South India) is subjected to seasonal changes in its physics and hydrochemistry variability and is a natural, dynamic situation (compared to a neighbourhood location of relative quietude) exhibiting features of catalysing the release of sediment bound phosphate depending on its utilisation, via bioturbation. The organic phosphate pool plays a crucial role, wherein it acts as a ‘inducer’ in the triggering mechanism of aquatic life in this coastal zone.     

Bacterial consumption of total and dissolved organic carbon in the Great Barrier Reef

Heterotrophic bacteria typically take up directly dissolved organic matter due to the small molecular size, although both particulate and dissolved organic matter have labile (easily consumed) compounds. Tropical coastal waters are important ecosystems because of their high productivity. However, few studies have determined bacterial cycling (i.e. carbon uptake by bacteria and allocation for bacterial biomass and respiration) of dissolved organic carbon in coastal tropical waters, and none has determined bacterial cycling of total and dissolved organic carbon simultaneously. In this study we followed bacterial biomass and production, and organic carbon changes over short-term (12 days) dark incubations with (total organic carbon, TOC) and without particulate organic carbon additions (dissolved organic carbon, DOC). The study was performed at three sites along the middle stretch of the Great Barrier Reef (GBR) during the dry and wet seasons. Our results show that the bacterial growth efficiency is low (0.1–11.5%) compared to other coastal tropical systems, and there were no differences in the carbon cycling between organic matter sources, seasons or locations. Nonetheless, more carbon was consumed in the TOC compared to the DOC incubations, although the proportion allocated to biomass and respiration was similar. This suggests that having more bioavailable substrate in the particulate form did not benefit bacteria. Overall, our study indicates that when comparing the obtained respiration rates with previously measured primary production rates, the GBR is a heterotrophic system. More detailed studies are required to fully explore the mechanisms used by bacteria to cycle TOC and DOC in tropical coastal waters.

     

The human impact in the German Bight: Eutrophication during three decades (1962–1991)

The human impact in the German Bight, in the form of anthropogenic eutrophication, has been documented by a 30-year time-series measurement near the island of Helgoland. Since 1962, the Biologische Anstalt Helgoland has measured inorganic nutrients and phytoplankton abundance from daily samples at Helgoland Roads, a position 60 km off the main source of eutrophication, the River Elbe. Since the early sixties, phosphate concentrations rose for about a decade, levelling off to about twice the former concentrations for another decade, and then decreasing (since 1982) as a result of phosphate-reducing measures. Nitrate concentrations, however, have only increased since 1980/81, following Elbe river flood events. In 1987, three times the former concentrations were reached. A decrease has been observed only since 1991. This different development of phosphorus and nitrogen eutrophication led to a shift of inorganic N/P-ratios in the German Bight. The phosphate increase was more pronounced in the late summer “regeneration mode” conditions, the nitrate increase in the winter months. The eutrophication is not restricted to the inner German Bight and coastal waters of a salinity of <33, but has also occurred in more saline waters at S>33 psu (practical salinity unit), as characteristic for the outer German Bight. In this more saline water, phosphate and nitrate maximum levels occurred three years later, compared with the average Helgoland data, which are more representative of the inner German Bight. It is suggested that suspended particulate organic matter, as a long-distance carrier of nutrients, might have caused this delayed eutrophication in the outer German Bight waters. While the human impact is obvious as to nutrient concentrations, it is less obvious in phytoplankton stock enhancement. A general increase in phytoplankton biomass (about 3–4 times) was found, but this was mainly due to unidentified nanoflagellates of unknown trophic state, and subject to methodological errors. The causal relationships of phytoplankton stocks and eutrophication are not clearly understood, as natural variability is large and hydrographical factors possibly dominate. Additional nutrient input by Elbe river floods did not always result in elevated phytoplankton stocks near Helgoland, while extended periods of vertical density stratification of the German Bight water caused large plankton blooms.     

Complex seasonal patterns of primary producers at the land-sea interface

Seasonal fluctuations of plant biomass and photosynthesis are key features of the Earth system because they drive variability of atmospheric CO₂, water and nutrient cycling, and food supply to consumers. There is no inventory of phytoplankton seasonal cycles in nearshore coastal ecosystems where forcings from ocean, land and atmosphere intersect. We compiled time series of phytoplankton biomass (chlorophyll a) from 114 estuaries, lagoons, inland seas, bays and shallow coastal waters around the world, and searched for seasonal patterns as common timing and amplitude of monthly variability. The data revealed a broad continuum of seasonal patterns, with large variability across and within ecosystems. This contrasts with annual cycles of terrestrial and oceanic primary producers for which seasonal fluctuations are recurrent and synchronous over large geographic regions. This finding bears on two fundamental ecological questions: (1) how do estuarine and coastal consumers adapt to an irregular and unpredictable food supply, and (2) how can we extract signals of climate change from phytoplankton observations in coastal ecosystems where local‐scale processes can mask responses to changing climate?     

A seasonal study of the distributions of surface state variables in Liverpool Bay. II. Nutrients

Distributions of dissolved inorganic nitrate, phosphate, and silicate observed in the surface waters of Liverpool Bay during a seasonal cycle of cruises in 1975 are presented. Winter distributions were influenced by the proportion of waters from various sources, their chemistry, and the mode of circulation, all of which were temporally variable. Non-winter distributions were additionally influenced by in situ biological activity. Principal component analysis provided a preliminary means of separating and assessing the relative importance of these factors and depicting the centres of biological activity during the non-winter cruises. Different circulation modes in January and December produced distributions with little visual similarity, and patchy nutrient distributions accompanied the spring and autumn phytoplankton blooms. Nitrogen-rich industrial and domestic effluents modified the nutrient availability ratios in the receiving waters so that silicon limited diatom growth over large areas of the Bay in spring. Nitrogen became ‘bio-limiting’ in summer and autumn. The direction of the surface residual circulation before the spring bloom is proposed as the major influence governing subsequent nutrient distributions and phytoplankton populations.     

Time scales and mechanisms of estuarine variability,a synthesis from studies of San Francisco Bay

This review of the preceding papers suggests that temporal variability in San Francisco Bay can be characterized by four time scales (hours, days-weeks, months, years) and associated with at least four mechanisms (variations in freshwater inflow, tides, wind, and exchange with coastal waters). The best understood component of temporal variability is the annual cycle, which is most obviously influenced by seasonal variations in freshwater inflow. The winter season of high river discharge is characterized by: large-scale redistribution of the salinity field (e.g. the upper estuary becomes a riverine system); enhanced density stratification and gravitational circulation with shortened residence times in the bay; decreased tissue concentrations of some contaminants (e.g. copper) in resident bivalves; increased estuarine inputs of river-borne materials such as dissolved inorganic nutrients (N, P, Si), suspended sediments, and humic materials; radical redistributions of pelagic organisms such as copepods and fish; low phutoplankton biomass and primary productivity in the upper estuary; and elimination of freshwater-intolerant species of macroalgae and benthic infauna from the upper estuary. Other mechanisms modulate this river-driven annual cycle: (1) wind speed is highly seasonal (strongest in summer) and causes seasonal variations in atmosphere-water column exchange of dissolved gases, resuspension, and the texture of surficial sediments; (2) seasonal variations in the coastal ocean (e.g. the spring-summer upwelling season) influence species composition of plankton and nutrient concentrations that are advected into the bay; and (3) the annual temperature cycle influences a few selected features (e.g. production and hatching of copepod resting eggs). Much of the interannual variability in San Francisco Bay is also correlated with freshwater inflow: wet years with persistently high river discharge are characterized by persistent winter-type conditions.Mechanisms of short-term variability are not as well understood, although some responses to storm events (pulses in residual currents from wind forcing, erosion of surficial sediments by wind waves, redistribution of fish populations) and the neap-spring tidal cycle (enhanced salinity stratification, gravitational circulation, and phytoplankton biomass during neap tides) have been quantified. In addition to these somewhat predictable features of variability are (1) largely unexplained episodic events (e.g. anomalous blooms of drift macroalgae), and (2) long-term trends directly attributable to human activities (e.g. introduction of exotic species that become permanent members of the biota).     

三门湾浮游动物的季节变动及微型浮游动物摄食影响

2002年8月、11月、2003年2月和5月,在三门湾进行了4个航次生物、化学和水文等专业综合调查。根据采集的浮游动物样品的分析鉴定及海上现场实验结果,对浮游动物的群落组成、生物量、丰度、多样性指数的分布和季节变动及其浮游动物对浮游植物的摄食影响进行研究。结果表明,三门湾浮游动物有67属,89种,16类浮游幼体,主要可划分为4个生态类群：以近岸低盐类群为主,其优势种为中华哲水蚤Calanus sinicus、真刺唇角水蚤Labidocera etwhaeta、捷氏歪水蚤Tortanus derjugini、太平洋纺锤水蚤Acartiapacifica、中华假磷虾Pseudeuphausia sinica和百陶箭虫Sagitta bedoti等。半咸水河口类群、暖水性外海类群和广布种相对较少。浮游动物生物量和丰度的平面分布趋势除了夏季有所差异外,其它季节基本一致。2月份和5月份,浮游动物生物量和丰度,从湾顶向湾口呈逐渐增加趋势;8月份,湾口区生物量最高,而丰度高值区出现在湾顶部;11月份,生物量和丰度的平面分布相对均匀。浮游动物种类多样性指数有明显的季节变化,其动态变化与浮游动物种数和丰度的变化一致。微型浮游动物对浮游植物存在摄食压力,且有季节变化,摄食率的变化在0．18．0．68d^-1,微型浮游动物的摄食率低于相同季节的浮游植物生长率。微型浮游动物对浮游植物摄食压力的变化范围为16．1％-49．1％d^-1,对初级生产力摄食压力的变化在58．3％-83．6％d^-1。11月份,微型浮游动物对浮游植物和初级生产力的摄食压力均出现最高值。     

Vulnerability and resilience of seagrasses to hurricane and runoff impacts along Florida’s west coast

Many climate change models predict increasing frequency and severity of tropical cyclones (hurricanes) in the Atlantic Ocean, Caribbean Sea, and Gulf of Mexico. To assess this potential threat to seagrass communities in Florida’s Big Bend region, we performed a habitat change analysis based on aerial seagrass surveys performed prior to, and after, the extremely active Atlantic cyclone seasons of 2004 and 2005. To provide a regional context for changes in the Big Bend region, we also compared impacts there with changes in three other West Florida estuaries. Our analysis showed that storm impacts on seagrasses varied along Florida’s west coast. Physical disturbance caused minor losses in parts of Charlotte Harbor and the Big Bend region. However, heavy rainfall in Florida and Georgia associated with Frances and Jeanne combined with winter rains to cause complete loss of 1,500 ha of seagrasses and thinning of another 1,700 ha in the vicinity of the Suwannee River mouth. In Tampa Bay, Sarasota Bay, and Charlotte Harbor, despite localized losses, total seagrass area actually increased between 2004 and 2006. On the other hand, Tampa Bay, Sarasota Bay, and Charlotte Harbor all showed significant, and more pronounced, declines in seagrass cover as the result of another major rainfall and runoff event: the 1997–1998 El Nino event. Our results indicate that light stress, likely caused by suspended sediments, phytoplankton blooms, and dissolved organic matter, resulted in seagrass losses extending up to 40 km from the mouth of the Suwannee River. We conclude that water quality impacts, especially if they are persistent, can be more damaging than physical impacts of moderate (Category 1–3) tropical cyclones. We also conclude that runoff-related impacts on seagrasses vary depending on the timing, volume, and persistence of storm runoff in relation to normal seasonal runoff patterns and seagrass growth in each estuary.     

东辽河流域河湖光学吸收特性的季节变化

吸收特性是水体重要的光学特性,也是建立水色反演分析模型的基本参数.本文利用2011年6、9、10月在二龙湖和2012年10月在东辽河的实测数据,通过测定水体营养水平和颗粒物、有色可溶性有机物(CDOM)等光学活性物质的吸收系数,分析了东辽河河湖的光学物质吸收特性和季节变化.结果表明: 二龙湖水体9月处于富营养状态,6月营养水平较低,10月则处于中营养水平;东辽河随采样点位置的差异呈现不同程度的富营养化.水体各组分的吸收系数均随营养水平的增加而逐渐增大.6、10月二龙湖水体总悬浮颗粒物吸收光谱与非色素颗粒物吸收光谱相似,在总吸收中占主导作用的是CDOM;9月总悬浮颗粒物吸收光谱则与浮游植物吸收光谱类似,浮游植物的吸收对总吸收的贡献率最大.6、9月二龙湖浮游植物吸收系数[a_ph(440)]与总磷(TP)呈正相关,6月二龙湖a_ph(440)与溶解有机碳(DOC)呈正相关,非藻类颗粒物吸收系数[a_d(440)]仅与叶绿素a浓度(Chl a)具有相关性.二龙湖(9月)与东辽河a_ph(675)均与Chl a、卡尔森指数(TLI)相关性较好.东辽河流域的河湖水体光学特性存在比较明显的差异:东辽河水体的浮游植物吸收光谱曲线展现出了两种不同类型,浮游植物、非藻类颗粒物、CDOM的贡献率随采样点的不同而相差较大;营养水平较高的9月二龙湖与东辽河水体浮游植物主导了水体光谱吸收,而在二龙湖营养水平较低的月份中(6、10月),颗粒物与CDOM共同主导水体光谱吸收;6月二龙湖与10月东辽河水体CDOM的组成为大分子颗粒物,而9与10月其组成更趋向于小分子颗粒物.     

Variability in factors causing light attenuation in Lake Victoria

1. The major optical components controlling the attenuation of photosynthetic available radiation in nearshore areas of Lake Victoria (Uganda and Kenya) were examined and their impact compared. It was found that chromophoric dissolved organic matter and tripton play a dominating role in many nearshore areas, indicating that the coastal areas of Lake Victoria cannot be considered as Case I waters.
2. Concentrations of chromophoric dissolved organic matter declined with distance from the coast in an exponential manner indicating dilution and degradation of terrestrial sources of organic matter rather than in situ production. The importance of tripton was found to follow a similar pattern, while the relative importance of phytoplankton biomass in overall attenuation of photosynthetic available radiation was found to increase with distance from the coast. A specific attenuation coefficient for phytoplankton biomass was determined (0.019 m² mg Chl a ⁻¹).
3. Using a light limitation approach based on carrying capacity, it was possible to map areas that are closer to being light limited. Light limitation appears to occur throughout most bays and some coastal areas receiving catchment waters. This spatial information, geographically referenced to bathymetric and catchment conditions, was utilized to understand the importance of environmental conditions in limiting phytoplankton biomass.     

Bacterial Growth Efficiency in a Tropical Estuary: Seasonal Variability Subsidized by Allochthonous Carbon

Bacterial growth efficiency (BGE) is a key factor in understanding bacterial influence on carbon flow in aquatic ecosystems. We report intra-annual variability in BGE, and bacteria-mediated carbon flow in the tropical Mandovi and Zuari estuaries (southwest India) and the adjoining coastal waters (Arabian Sea). BGE ranged from 3% to 61% and showed clear temporal variability with significantly (ANOVA, p < 0.01) higher values in the estuaries (mean, 28 ± 14%) than coastal waters (mean, 12 ± 6%). The greater variability of BGE in the estuaries than coastal waters suggest some systematic response to nutrient composition and the variability of dissolved organic matter pools, as BGE was governed by bacterial secondary production (BP). Monsoonal rains and its accompanied changes brought significant variability in BGE and bacterial productivity/primary productivity (BP/PP) ratio when compared to nonmonsoon seasons in the estuaries and coastal waters. High BP/PP ratio (>1) together with high carbon flux through bacteria (>100% of primary productivity) in the estuarine and coastal waters suggests that bacterioplankton consumed dissolved organic carbon in excess of the amount produced in situ by phytoplankton of this region, which led to the mismatch between primary production of carbon and amount of carbon consumed by bacteria. Despite the two systems being subsidized by allochthonous inputs, the low BGE in the coastal waters may be attributable to the nature and time interval in the supply of allochthonous carbon.     

Environmental impact of a salmonid farm on a well flushed marine site: I. Current and water quality

The impact of a common brown trout farm on water quality was investigated at a site well flushed by lapping and tidal current. Dissolved oxygen, suspended matter, phosphate, nitrite and nitrate were seemingly not affected by the farm. High concentrations of ammonia were clearly observed close to the raft, but rapidly vanished a short distance from it, when the fish biomass was at least 576 tonnes. At that time the nitrogen loadings from the farm were estimated at 101 kg/day. On the site, phytoplankton was similar to that of the nearby French coastal area of the English Channel, concerning generic composition (extensively dominated by diatoms) and the seasonal succession of genera. The farm did not cause any increase of faecal coliform bacteria. At this stock level, the dispersal characteristics of the farm site greatly reduced its apparent impact on the water quality.     

Statistical analysis of phytoplankton biomass in coastal waters: Case study of the Wadden Sea near Lauwersoog (The Netherlands) from 2000 to 2009

As the most important indicator of the coastal ecosystem, various studies on phytoplankton have frequently been proposed in coastal waters. In this study, the statistical analysis of phytoplankton biomass (in terms of chlorophyll a) is performed in the Wadden Sea near Lauwersoog (53.4167° N, 6.2000° E), The Netherlands, determined by the 10-year's historical dataset from 2000 through 2009. Boxplot analysis is introduced to give insight in the seasonal variations of phytoplankton biomass and physical–chemical conditions. Annually, a big difference is found in chlorophyll a, showing that the maximum values occur in the months of April and July, and the minimum values occur in winter (December, January, and February). The phytoplankton biomass is positively correlated with the physical conditions (salinity, light intensity, and water temperature), and is negatively correlated with the nutrients (nitrate, ammonium, and silicate). Factor analysis distinguishes the driving variables that represent most of the total variance using three extraction methods (principal component analysis, unweighted least squares, and maximum likelihood). Nitrate dominates the main activity in the first rotated component/factor, and ammonium contributes the most in the second rotated component/factor. The findings of this study are significant for understanding the roles of the environmental conditions upon the phytoplankton variability, and for reducing the overlapping information to characterize the driving variables.     

Biochemical markers of commercial fish adaptation in estuaries of Peter the Great Bay (the Sea of Japan)

The results of the comparative study of molecular biomarkers of oxidative stress and biotransformation of some representatives of the families Cyprinidae and Mugilidae from estuaries of rivers which enter Peter the Great Bay are presented. The haarder mullet Liza haematocheila is used as an indicator species for monitoring the degree of environmental pollution. The basic values of biochemical parameters in different organs of haarder mullet and their seasonal variations have been determined. A significant increase in the concentration of glutathione and activity of glutathione-S-transferase in fish from the Razdolnaya River has been found on the basis of the comparison of the functional state of haarder mullet in estuaries of the Razdolnaya and Amba rivers, which differ in the degree of anthropogenic load.     

Nutrient limitation of phytoplankton growth in Waquoit Bay, Massachusetts, USA: a nutrient enrichment study

We conducted nutrient enrichment experiments and field sampling to address three questions: (1) is there nutrient limitation of phytoplankton accumulation within an estuary whose waters are exposed to relatively high nitrogen loading rates, (2) where in the salinity gradient from fresh to seawater (0 to 32‰) is there a shift from phosphorus to nitrogen limitation of phytoplankton accumulation, and (3) is there a seasonal shift in limiting function of phosphorus and nitrogen anywhere in the estuarine gradient. Nitrogen and phosphorus enrichment experiments in the Childs River, an estuary of Waquoit Bay, Massachusetts, USA, showed that the accumulation of phytoplankton biomass in brackish and saline water was limited by supply of nitrate during warm months. The effects of enrichment were less evident in fresh water, with short-lived responses to phosphate enrichment. There was no specific point along the salinity gradient where there was a shift from phosphorus- to nitrogen-limited phytoplankton accumulation; rather, the relative importance of nitrogen and phosphorus changed along the salinity gradient in the estuary and with season of the year. There was no response to nutrient additions during the colder months, suggesting that some seasonally-varying factor, such as light, temperature or a physiological mechanism, restricted phytoplankton accumulation during months other than May-Aug. There was only slight evidence of a seasonal shift between nitrogen- and phosphorus-limitation of chlorophyll accumulation. Phytoplankton populations in nutrient-rich estuaries with short flushing times grow fast, but at the same time the cells may be advected out of the estuaries while still rapidly dividing, thereby providing an important subsidy to production in nearby deeper waters. This revised version was published online in August 2006 with corrections to the Cover Date.