Nutrients,light and primary production by phytoplankton and microphytobenthos in the eutrophic,turbid Westerschelde estuary (The Netherlands)

Abiotic factors and primary production by phytoplankton and microphytobenthos was studied in the turbid Westeschelde estuary. Because of the high turbidity and high nutrient concentrations primary production by phytoplankton is light-limited. In the inner and central parts of the estuary maximum rates of primary production were therefore measured during the summer, whereas in the more marine part spring and autumn bloom were observed. Organic loading is high, causing near anaerobic conditions upstream in the river Schelde. Because of this there were no important phytoplankton grazers in this part of the estuary and hence the grazing pressure on phytoplankton was minimal. As this reduced losses, biomass is maximal in the river Schelde, despite the very low growth rates.On a number of occasions, primary production by benthic micro-algae on intertidal flats was studied. Comparison of their rates of primary production to phytoplankton production in the same period led to the conclusion that the contribution to total primary production by benthic algae was small. The main reason for this is that the photosynthetic activity declines rapidly after the flats emerged from the water. It is argued that CO₂-limitation could only be partially responsible for the noticed decrease in activity.     

Changes in phytoplankton biomass and primary production between 1991 and 2001 in the Westerschelde estuary (Belgium/The Netherlands)

The Westerschelde estuary is a very polluted and turbid estuary, but the last decade the waterquality improved. Dredging activity also increased in 1997 to allow bigger ships to enter the port of Antwerpen. This could potentially decrease the light conditions for the phytoplankton. Because of all these recent changes in the estuary we studied primary productivity in 2001 and compared it to values in 1991. The results show that due to a decrease in discharge in particulate and dissolved organic carbon the oxygen concentrations in general have increased in the upstream region, although in spring and summer low oxygen concentrations (10–30% saturation) can still be found. Phosphate and ammonia concentrations have decreased and the zone of nitrification which was very large in 1991 has become very small and is now located in the uppermost upstream region of the estuary. Si-concentrations have remained the same. All nutrient concentrations are still high enough not to limit phytoplankton growth. Turbidity remained unaltered as a result of the dredging works, and as a result phytoplankton biomass in most of the estuary did not show a decrease, although there were signs that in the upstream region phytoplankton biomass decreased, possible caused by increased grazing pressure. The relationship between phytoplankton biomass and primary productivity did not change, and from the data it can be concluded that the dredging activity will not influence the gross and net primary productivity of the phytoplankton.     

Inter-annual,seasonal and spatial variability in nutrient limitation of phytoplankton production in a river impoundment

We characterize seasonal and spatial patterns in phytoplankton abundance, production and nutrient limitation in a mesotrophic river impoundment located in the southeastern United States to assess variation arising from inter-annual differences in watershed inputs. Short-term (48 h) in situ nutrient addition experiments were conducted between May and October at three sites located along the longitudinal axis of the lake. Nutrient limitation was detected in 12 of the 18 experiments conducted over 2 years. Phytoplankton responded to additions of phosphorus alone although highest chlorophyll concentrations were observed in enclosures receiving combined (P and N) additions. Growth responses were greatest at downstream sites and in late summer suggesting that those populations experience more severe nutrient limitation. Interannual variation in nutrient limitation and primary production corresponded to differences in the timing of hydrologic inputs. Above average rainfall and discharge in late-summer (July–October) of 1996 coincided with higher in-lake nutrient concentrations, increased production, and minimal nutrient limitation. During the same period in 1995, discharge was lower, nutrient concentrations were lower, and nutrient limitation of phytoplankton production was more pronounced. Our results suggest that nutrient limitation is common in this river impoundment but that modest inter-annual variability in the timing of hydrologic inputs can substantially influence seasonal and spatial patterns.     

Linking population dynamics models with empirically derived models through phytoplankton primary production

There are alternative methods for estimation of phytoplankton primary production (PP) that are fundamentally different in the calculation approach. The process‐oriented PP model is a mechanistic, empirically derived method based on the photosynthesis–light relationships. The population dynamics‐based PP calculation, which is a synthetic method, provides a production estimate based on population dynamics of phytoplankton. These alternative methods were here compared with regard to production estimates and linked to enhance the performance of the existing models of population dynamics applied to a wide variety of lakes worldwide in terms of morphometry, nutrient status, and light environments. Estimates of PP were shown to be sensitive to changes in phytoplankton sinking and zooplankton grazing rates in both methods. Production estimates in the process‐oriented PP model were also sensitive to light‐associated parameters such as day length. Although the production estimated from the population dynamics‐based PP calculation tended to be lower than that from the process‐oriented PP model irrespective of lake morphometry, production estimates calculated from both methods with standard parameterization were comparable when production was estimated on an annual timescale. However, it was also shown that the alternative methods could produce different production estimates when estimated on shorter timescales such as cyanobacterial blooms in summer. Cyanobacteria with low mortality due to grazing and sinking losses have been considered as trophic bottlenecks, but there is increasing evidence that their mortality is, to a considerable extent, due to parasitic pathogens. In the case of cyanobacterial blooms, an addition of parasite‐related loss term (19%–33% of standing stock) resulted in a resolution of the difference in production estimates between the methods. These analyses theoretically support the critical role of parasitism and resolve the bottleneck problem in aquatic ecosystem metabolism.     

Physical and chemical characters,phytoplankton and primary production of Ezequiel Ramos Mexía Reservoir (Argentina)

We describe the distribution in space and time of a series of physical and chemical variables, phyto-plankton, and primary production in Ezequiel Ramos Mexía Reservoir (Argentina). Its waters are soft, poor in nutrients and with a low transparency that greatly depresses primary production. Phytoplankton data indicate the presence of 79 taxa with Bacillariophyceae, Cyanophyta and Chlorophyta alternatively dominant. Chlorophyll a was always low and never exceeded 3 mg m⁻³. Based on these results, the trophic status of this ecosystem is discussed.     

The primary production of phytoplankton in Lake Vechten

The primary production of the phytoplankton of Lake Vechten (The Netherlands) (area, 4.7 ha; mean depth, 6 m), an unpolluted and stratified sandpit was investigated from 1969 to 1980 (except in 1971, 1975 and 1976) by the in situ ¹⁴C-technique. Other data collected include: solar radiation, transparency, oxygen and thermal structure. In winter and spring diatoms, Cryptophyceae and Chlorococcales were important algal groups, while in summer Dinophyceae and Chrysophyceae were important. The chlorophyll-a concentration was compared to the cellular biovolumes (= fresh weight) of the most abundant phytoplankton species. The primary production maxima occurred in winter, spring and during the summer stratification. The vertical profiles of photosynthesis exhibit light inhibition at surface to a maximum of 4 m. The maximum of zooplankton grazing in May–June caused a sharp decrease in the phytoplankton biomass and seston concentration accompanied by the highest transparency (clear water phase).The values for cellular C-fixation range from 10 to 1307 mg C · M^–2 · day^–1 (annual mean of 280 mg C · m^–2 · day^–1). High dark fixation (up to 100%) was encountered in the metalimnion and hypolimnion from July to October together with peaks of ¹⁴C-fixation due to a crowding of phytoplankton and phototrophic anoxic bacteria. Extracellular excretion by phytoplankton, investigated in 1977 to 1979, was 15% of the annual mean of the total C-fixation. The photosynthetic efficiency, turnover rates, and activity coefficients were low, particularly in the summer months when Ceratium hirundinella was predominant. The seasonal variations were controlled mainly by solar radiation and probably phosphate, the former being more important in the non-stratification period and the latter during the stratification period.     

Distinction between light-mediated and light-independent variations in phytoplankton production rates

An array of factors simultaneously controls phytoplankton photosynthesis and hence the primary production process. Because their relative importance shifts both with depth and with season, the significance of individual factors cannot be resolved by in situ incubations, even if all relevant environmental and biotic variables are measured.Here a procedure is described by which in addition to in situ measurements, photosynthesis is simultaneously assessed in identical subsamples under constant temperature (10 °C) and light (0.66 mol m^–2 h^–1 PAR conditions, in vitro). By calculating photosynthesis per unit of chlorophyll, effects of shifting biomass on photosynthesis can be eliminated but seasonal variations of light-saturated photosynthesis generated by temperature, and vertical changes of light-requirements (e.g. by light-shade adaptation) remain obscure. Quotients of in situ photosynthetic rates divided by in vitro rates allow the quantification of light-mediated changes. Provided that photosynthesis measured in vitro is light-saturated, quotients in situ: in vitro rates should never exceed unity. They are a measure for the degree of light-limitation. In vitro rates normalized to chlorophyll give information on temporal changes caused by variations in photosynthetic capacity. In Lake Constance, mean cell size appears to control light-saturated assimilation numbers.     

瓯江口春季营养盐、浮游植物和浮游动物的分布

研究2007年4月瓯江口海域(27°38′～28°02′N、120°50′～121°14′E)浮游生物空间分布特征,分析这一分布特征与营养盐和其他水文要素之间的联系.结果表明,悬浮物浓度、DIP和DIN分布特征均是近河口最高,由近河口向外数量逐渐减少.在灵昆岛南侧和东南侧近口门水域,是DIP和DIN的高值中心,但该水域高浓度悬沙使水体透光率较低,不利于浮游植物生长.在口门外侧海域,悬沙浓度已经明显降低,因而是浮游植物高密度区域.浮游植物和浮游动物丰度分布趋势基本相同,由内海向外海数量逐渐递减;其中,大型浮游动物丰度和浮游植物丰度,小型浮游动物丰度和浮游植物丰度之间显著地呈线性正相关关系.这一特征的形成,主要由瓯江口营养盐和悬浮物分布特征决定,浮游动物与浮游植物空间分布的一致性,很好地反映出浮游动物对浮游植物有效的下行控制,从而使水域生态系统在此季节保持稳定.     

Seasonal variation in phytoplankton primary production in relation to light and nutrients in Lake Awasa,Ethiopia

The biomass and primary production of phytoplankton in Lake Awasa, Ethiopia was measured over a 14 month period, November 1983 to March 1985. The lake had a mean phytoplankton biomass of 34 mg chl a m^–3 (n = 14). The seasonal variation in phytoplankton biomass of the euphotic zone (mg chl a m^–2 h^–1) was muted with a CV (standard deviation/mean) of 31%. The vertical distribution of photosynthetic activity was of a typical pattern for phytoplankton with light inhibition on all but overcast days. The maximum specific rates of photosynthesis or photosynthetic capacity (Ø_max) for the lake approached 19 mg O₂ (mg chl a)^–1 h^–1, with high values during periods of low phytoplankton biomass. Areal rates of photosynthesis ranged between 0.30 to 0.73 g O₂ m^–2 h^–1 and 3.3 to 7.8 g O₂ m^–2 d^–1. The efficiency of utilisation of PhAR incident on the lake surface varied from 2.4 to 4.1 mmol E^–1 with the highest efficiency observed corresponding to the lowest surface radiation. Calculated on a caloric basis, the efficiency ranged between 1.7 and 2.9%. The temporal pattern of primary production by phytoplankton showed limited variability (CV = 21 %).     

Effects of nutrient and light limitation on the biochemical composition of phytoplankton

Three marine phytoplankters (Isochrysis galbana, Chaetoceros calcitrans andThalassiosira pseudonana), commonly used in the culture of bivalve larvae, were grown in batch or semi-continuous cultures. Changes in protein, carbohydrate, lipid and some fatty acids were measured as growth became limited by nitrogen, silicon, phosphorus or light. Under N starvation (2 d) the % lipid remained relatively constant, while% carbohydrate increased and% protein decreased in all 3 species compared to cells growing under no nutrient limitation. Under Si starvation (6 h) there was no change in lipid, protein or carbohydrates. The amount of two fatty acids, 20 : 53 and 22 : 63 remained relatively constant under N, P and Si starvation, exept for a sharp drop in the cells of P-starvedT. pseudonana. However, there were pronounced species differences withI. galbana containing significantly less 20 : 5 3 thanC. calcitrans orT. pseudonana. Under light limitation the amount of lipid per cell showed no consistent trend over a range of irradiances for all 3 species. The amount of N per cell (an index of protein content) as a function of irradiance, was relatively constant forI. galbana andT. pseudonana, while the amount of N per cell was lower under low irradiances forC. calcitrans. These examples of changes in protein, carbohydrate, lipid and certain fatty acids under nutrient (N, Si or P) or light limitation, emphasize the importance of knowing the phase (e.g. logarithmic vs stationary) of the growth curve in batch cultures, since the nutritional value of the phytoplankters could change as cultures become dense and growth is terminated due to nutrient or light limitation.Presented at the XIIIth International Seaweed Symposium, University of British Columbia, Vancouver, Canada, August 1989.     

Assessment of primary production in the White Sea

The literature and original data on the primary production of phytoplankton in the White Sea are analyzed. By this parameter, the White Sea is significantly inferior only to the Chukchi Sea; it is similar to the Barents Sea, and exceeds other Russian Arctic seas by two to three times (the Kara Sea, Laptev Sea, and East Siberian Sea).     

Primary production of phytoplankton in Lake Valencia (Venezuela)

Lake Valencia is heavily polluted by waste water of domestic, agricultural and industrial origin. The high organic load may have produced important changes in the limnological properties. Cyanobacteria dominated in numbers and biomass (over 90% throughout the year). Chlorophyll-a content averaged 37.7 + 15 μg · 1⁻¹. Maximum concentrations of 50–80 μg · 1⁻¹ were found near the inflows affected by organically polluted affluents. There has been a 50% reduction in the euphotic zone in only 13 years. The maximum rate of gross photosynthesis per hour at light saturation was determined within the uppermost 1-meter layer. The highest value was 16,290 mg O₂ · m⁻³ · h⁻¹. Lake Valencia is among the most productive lakes in the world, with areal net photosynthesis averaging 7.5 g C · m⁻² · d⁻¹.     

Aspects of winter primary production in the upstream section of Saguenay Fjord

From August 1985 to May 1986, experiments on primary production were carried out bi-weekly in the superficial waters (1–10 m) at one representative station (maximum depth of 260 m) located at the head of the Saguenay Fjord. During the winter season (January–March), the microalgal cells are abundant but their productivity is low. At the beginning of the snow-ice cover (January) the cells are mainly concentrated in the ice-cover interface and in the surface waters whereas in February–March, the cells are much more abundant in the ice. Results showed that the physico-chemical conditions are unfavourable for high productivity. The quantity and the quality of light under the ice cover are not the only factors limiting the photosynthetic activity of epontic community. The ice interior and epontic communities during the winter are mostly composed of the two euryhaline speciesAsterionella formosa andTabellaria fenestrata.     

Seasonal change in the proportion of bacterial and phytoplankton production along a salinity gradient in a shallow estuary

We intended to evaluate the relative contribution of primary production versus allochthonous carbon in the production of bacterial biomass in a mesotrophic estuary. Different spatial and temporal ranges were observed in the values of bacterioplankton biomass (31–273 g C l^–1) and production (0.1–16.0 g C l^–1 h^–1, 1.5–36.8 mg C m^–2 h^–1) as well as in phytoplankton abundance (50–1700 g C l^–1) and primary production (0.1–512.9 g C l^–1 h^–1, 1.5–512.9 mg C m^–2 h^–1). Bacterial specific growth rate (0.10–1.68 d^–1) during the year did not fluctuate as much as phytoplankton specific growth rate (0.02–0.74 d^–1). Along the salinity gradient and towards the inner estuary, bacterio- and phytoplankton biomass and production increased steadily both in the warm and cold seasons. The maximum geographical increase observed in these variables was 12 times more for the bacterial community and 8 times more for the phytoplankton community. The warm to cold season ratios of the biological variables varied geographically and according to these variables. The increase at the warm season achieved its maximum in the biomass production, particularly in the marine zone and at high tide (20 and 112 times higher in bacterial and phytoplankton production, respectively). The seasonal variation in specific growth rate was most noticeable in phytoplankton, with seasonal ratios of 3–26. The bacterial community of the marine zone responded positively – generating seasonal ratios of 1–13 in bacterial specific growth rate – to the strong warm season increment in phytoplankton growth rate in this zone. In the brackish water zone where even during the warm season allochthonous carbon accounted for 41% (on average) of the bacterial carbon demand, the seasonal ratio of bacterial specific growth rate varied from about 1 to 2. During the warm season, an average of 21% of the primary production was potentially sufficient to support the whole bacterial production. During the cold months, however, the total primary production would be either required or even insufficient to support bacterial production. The estuary turned then into a mostly heterotrophic system. However, the calculated annual production of biomass by bacterio- and phytoplankton in the whole ecosystem showed that auto- and heterotrophic production was balanced in this estuary.     

Relations between production and biomass of phytoplankton in four Swedish lakes of different trophic status and humic content

Production and biomass values from phytoplankton populations in four different Swedish lakes were analysed. The production in all lakes was directly proportional to biomass during homothermal periods. When the lakes were stratified there was a strong negative relation between specific growth rate and biomass. The data fitted to a logistic density dependent growth equation of the form: dB/ dt = µ_mB(1-B · K^–1) where B is the biomass, µ_m the maximum specific growth rate and K the carrying capacity. The equation was used to derive the parameters µ · µ_m ^–1 and carrying capacity (the maximum possible biomass). These parameters were then discussed in relation to light climate, phosphorus concentration and humic content.     

The influence of mesozooplankton on phytoplankton nutrient limitation: a mesocosm study with northeast Atlantic plankton

We used marine phytoplankton from mesocosms seeded with different zooplankton densities to study the impact of mesozooplankton on phytoplankton nutrient limitation. After 7 d of grazing (copepod mesocosms) or 9 d (appendicularian mesocosms) phytoplankton nutrient limitation was studied by enrichment bioassays. After removal of mesozooplankton, bioassay bottles received either no nutrients, phosphorus or nitrogen alone, or a combination of nitrogen and phosphorus and were incubated for 2 d. Phytoplankton reproductive rates in the bottles without nutrient addition were calculated after correction for grazing by ciliates and indicated increasing nitrogen limitation with increasing copepod abundance. No nutrient limitation was found in the appendicularian mesocosms. The increase of nutrient limitation with increasing copepod density seems to be mainly the result of a trophic cascade effect: Copepods released nanoplankton from ciliate grazing pressure, and thereby enhanced nitrogen exhaustion by nanophytoplankton and reduced nitrogen excretion by ciliates. Nitrogen sequestration in copepod biomass, the mechanism predicted by the ecological stoichiometry theory, seems to have been a weaker effect because there was only little copepod growth during the experiment.     

Surface water CO2 concentration influences phytoplankton production but not community composition across boreal lakes

Recent experimental evidence suggests that changes in the partial pressure of CO₂ (pCO₂), in concert with nutrient fertilisation, may result in increased primary production and shifted phytoplankton community composition that favours species lacking adaptations to low CO₂ environments. It is not clear whether these results apply in ambient freshwaters, which are already often supersaturated in CO₂, and where phytoplankton structure and activity are under complex control of diverse local and regional factors. Here, we use a large‐scale comparative study of 69 boreal lakes to explore the influence of existing CO₂ gradients (c. 50–2300 μatm) on phytoplankton community composition and biomass production. While community composition did not respond to pCO₂ gradients, gross primary production was enhanced, but only in lakes already supersaturated in CO₂, demonstrating that environmental context is key in determining pCO₂–phytoplankton interactions. We further argue that increased atmospheric CO₂ is unlikely to influence phytoplanktonic composition and production in northern lakes.     

Effects of nutrient addition on phytoplankton in Lake Nakanuma,Japan I. Changes in biochemical components

Effects of nutrient conditions on biochemical components (sugars and amino acids) of phytoplankton were examined in Lake Nakanuma in Japan. Phosphate, ammonium and silicate were added to water samples collected at 0 m, which were incubated for 15 days in situ. Chlorophyll a in phosphate-added samples increased much more than that in other samples. Total amino acids and total neutral sugars in phosphate-added samples also increased more than those in other samples. The increase of total amino acids and chlorophyll began faster than that of total neutral sugars during the first 5 days of incubation. Total neutral sugars in the phosphate-added samples increased rapidly after 8 days. The composition of amino acids did not change so much. However, the composition of neutral sugars changed according to the different nutrient addition. Phosphate-added samples changed greatly during the incubation. These changes were explained at least partly by changes in nutrient conditions. Addition of limiting nutrients decreased glucose content, whereas depletion of nutrients increased the content. This study indicates that measurements of the biochemical components contribute to the analysis of effects of nutrients on phytoplankton in natural waters.     

2006年冬季南海北部浮游植物生物量和初级生产力及其环境调控

通过2006年2月在南海北部现场调查所获得的数据分析研究了海域的浮游植物现存量和生产力,结果表明冬季浮游植物的分布特征与东北季风引起的环流场的变化关系密切。调查海域水柱平均Chla浓度的变化范围为0.03～1.21mg·m-3（平均（0.33±0.33）mg·m^-3）,高值区出现在广东沿岸及海南岛东部附近海域;初级生产力的分布范围为41.3～1040.0mgC·m^-2d^-1。由于东北季风引起近岸水体混合剧烈,不利于浮游植物生长,因此虽然沿岸带Chla浓度（（0.53±0.50）mg·m^-3）较高,但初级生产力却是最低的,只有41.3mgC·m^-2d^-1,同时冬季反气旋涡强度下降和底层富营养水的涌升,营养盐充足,因此开阔海的Chla浓度（（0.31±0.30）mg·m^-3）和初级生产力（（631.3±578.0）mgC·m^-·2d^-1）均高。浮游植物粒度级份测定表明,Pico级份对冬季南海北部浮游植物现存量和生产力的贡献很大,特别是在开阔海区,分别占47%和66%。     

珠江口及毗邻海域营养盐对浮游植物生长的影响

基于2006年7月(夏季),10月(秋季)和2007年3月(春季)的现场调查数据,对珠江口及毗邻海域中的营养盐和叶绿素a等环境生态因子的时空分布特性进行了对比分析,研究了氮磷比与叶绿素a含量和种群多样性之间的联系,探讨了该海域营养盐对于浮游植物生长的影响。结果表明:(1)研究海域营养盐表现出较强的季节和空间差异性,总氮(TN)和总磷(TP)浓度均值春季(1.545 mg/L、0.056 mg/L)和夏季(1.570 mg/L、0.058 mg/L)均大于秋季(1.442 mg/L、0.034 mg/L),且春夏季浓度空间差异更明显。(2)调查期间海域营养盐含量超标现象突出,夏季尤为明显。无机氮(DIN)总体均值0.99 mg/L,超四类海水标准限值1倍,活性磷酸盐(PO4-P)总体均值0.021 mg/L,DIN∶PO4-P平均值为130;叶绿素a浓度与营养盐、p H、温度有较显著的相关性。(3)叶绿素a浓度较高的站位,具有较高的DIN∶PO4-P值,但浮游植物多样性指数偏低,优势种明显,主要为中肋骨条藻。氮磷比的改变会影响不同生长特性的浮游植物间的竞争和种群结构的改变;今后海洋污染治理中,在控制氮、磷污染时要注意氮磷比的改变可能造成的浮游生态影响。