The contribution of phytoplankton productivity in turbid freshwaters to their trophic status

Certain factors influencing phytoplankton productivity are accentuated in turbid waters. They include mixing, spectral quality shifts, scattering, light fluctuations, and overall light attenuation. Measurements of productivity is influenced by the presence of inorganic turbidity. Together with the above factors high turbidity causes difficulties to assess and model phytoplankton productivity. Estimations of ^B, P_m ^B, I_k and _m only reflect on the physiological condition of the phytoplankton, which differs little between water types of temperate regions. Measurement of integral vertical productivity, efficiency and fractional absorption by the phytoplankton of light energy conversion, however, are greatly influenced by inorganic turbidity. Because of high ratios of mixing to euphotic depth, the critical mixing depth is one of the most important factors influencing overall productivity in turbid waters.     

河口浮游植物生态学研究进展

综述了河口浮游植物种类组成、时空分布、初级生态及其影响因素等方面的主要研究进展,同时,对河口浮游植物在水环境监测中的作用以及河口浮游植物多样性与边缘效应进行了初步探讨。研究表明,通常河口区重要的浮游植物有硅藻、甲藻等,微型、超微型浮游植物在河口生态系统中占有重要地位。河口浮游植物种类组成、初级生产的时空变化明显,并受到光、温度、营养盐、动物摄食以及径流等因素的影响。     

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.     

Response of Phytoplankton Photophysiology to Varying Environmental Conditions in the Sub-Antarctic and Polar Frontal Zone

Climate-driven changes are expected to alter the hydrography of the Sub-Antarctic Zone (SAZ) and Polar Frontal Zone (PFZ) south of Australia, in which distinct regional environments are believed to be responsible for the differences in phytoplankton biomass in these regions. Here, we report how the dynamic influences of light, iron and temperature, which are responsible for the photophysiological differences between phytoplankton in the SAZ and PFZ, contribute to the biomass differences in these regions. High effective photochemical efficiency of photosystem II (/ 0.4), maximum photosynthesis rate (), light-saturation intensity (), maximum rate of photosynthetic electron transport (1/), and low photoprotective pigment concentrations observed in the SAZ correspond to high chlorophyll and iron concentrations. In contrast, phytoplankton in the PFZ exhibits low / ( 0.2) and high concentrations of photoprotective pigments under low light environment. Strong negative relationships between iron, temperature, and photoprotective pigments demonstrate that cells were producing more photoprotective pigments under low temperature and iron conditions, and are responsible for the low biomass and low productivity measured in the PFZ. As warming and enhanced iron input is expected in this region, this could probably increase phytoplankton photosynthesis in this region. However, complex interactions between the biogeochemical processes (e.g. stratification caused by warming could prevent mixing of nutrients), which control phytoplankton biomass and productivity, remain uncertain.     

Effect of phytoplankton size diversity on primary productivity in the North Pacific: trait distributions under environmental variability

While most biodiversity and ecosystem functioning (BEF) studies have found positive effects of species richness on productivity, it remain unclear whether similar patterns hold for marine phytoplankton with high local richness. We use the continuous trait‐based modelling approach, which assumes infinite richness and represents diversity in terms of the variance of the size distribution, to investigate the effects of phytoplankton size diversity on productivity in a three‐dimensional ocean circulation model driven by realistic physics forcing. We find a slightly negative effect of size diversity on primary production, which we attribute to several factors including functional trait‐environment interactions, flexible stoichiometry and the saturation of productivity at low diversity levels. The benefits of trait optimisation, whereby narrow size distributions enhance productivity under relatively stable conditions, tend to dominate over those of adaptive capacity, whereby greater diversity enhances the ability of the community to respond to environmental variability.     

Primary and new production in the deep Canada Basin during summer 2002

The NOAA Ocean Exploration program provided the opportunity to measure the carbon and nitrogen productivity across the Canada Basin. This research examined the major environmental factors limiting the levels of primary production and possible future climate change on the ecosystems. The vertical distributions of the carbon and nitrogen uptakes of phytoplankton had similar patterns as their respective biomass concentrations which were low at the surface and highest in the chlorophyll-maximum layer. The annual carbon and new production rates of phytoplankton in the Canada Basin were about 5 and 1 g C m^–2, respectively. Nutrients were determined to be a main limiting factor at the surface, whereas light may be a major factor limiting phytoplankton productivity in the chlorophyll-maximum layer for open waters. The bottom surface of the ice has a low specific uptake and productivity of phytoplankton, indicating that photosynthetic activity might be controlled by both light and nutrients.     

Phytoplankton productivity and its response to higher light levels in the Canada Basin

Phytoplankton productivity in the Canada Basin was measured in the late summer season, from mid-September to mid-October 2009, using a ¹³C–¹⁵N dual tracer technique. To understand potential production changes associated with sea ice melting in the Arctic Ocean, we examined the effects of light enhancement and nitrate enrichment on the carbon productivity of phytoplankton from the chlorophyll a maximum layer. The daily carbon productivity in the Canada Basin in 2009 was very low, with a mean of 4.1 mg C m⁻² (SD = 3.6 mg C m⁻²), compared with those reported in previous studies in the region. Among several explanations, the most plausible reason for the large difference in carbon productivity between this and the previous studies was strong seasonal variation in biomass and photosynthetic rate of the phytoplankton in the study region. Based on our results from light enhancement and nitrate enrichment experiments, we found that carbon productivity of phytoplankton in the chlorophyll a maximum layer could be stimulated by increased light condition rather than nitrate addition. Thus, potentially increasing light availability from current and ongoing decreases in the sea ice cover could increase the carbon production of the phytoplankton in the chlorophyll a maximum layer and produce a well-developed maximum layer at a deeper depth in the Canada Basin.     

Underwater light field and phytoplankton absorbance in different surface water masses of the Atlantic sector of the Southern Ocean

Spectral water transparency and phytoplankton light absorbance were studied in the Atlantic sector of the Southern Ocean during the Southern Ocean JGOFS ANT XIII/2 cruise in early austral summer 1995/1996. The study area comprised three zones, which differed markedly with respect to their hydrographic and planktological characteristics: the Antarctic Polar Frontal Zone with adiatom bloom, the Antarctic Circumpolar Current outside frontal systems with phytoplankton-poor water and a higher flagellate abundance than in the other two areas, and the marginal ice zone with a Phaeocystis bloom. The influence of phytoplankton on spectral water transparency was assessed by two independent procedures: the pigment-specific beam absorption coefficient, a_J*[5], at all stations, as estimated by spectroscopy of in vivo light absorption of plankton on glass fibre filters, and the pigment-specific light attenuation, (k_c[5]), as derived by regression analysis of spectral in situ vertical light attenuation coefficients in the sea against concomitant pigment concentrations. Values of a_J*[5] and vertical profiles of light attenuation by phytoplankton exhibited regional differences that corresponded with the three zones from which samples had been collected. These differences can be related to the specific characteristics of the three zones with respect to cell size distribution, pigment composition and biomass. The observed variations in a_J*[5] values should be considered when oceanic primary production is to be estimated by biooptical modelling.     

Carbon fixation and carbon availability in marine phytoplankton

It is widely believed that inorganic C does not limit the rate of short-term photosynthesis, the net productivity, or the maximum biomass, of marine phytoplankton. This lack of inorganic C restriction is less widely believed to hold for phytoplankton in many low alkalinity freshwaters or for seaweed in nutrient-enriched rock pools. These views are examined in the context of the physical chemistry of the inorganic C system in natural waters and of the ways in which various taxa of phytoplankton deal with inorganic C and discriminate between ¹²C and ¹³C. Using this information to interpret data obtained in the ocean or in freshwater suggests that short-term photosynthesis, production rate, and achieved biomass, of phytoplankton are rarely limited by inorganic C supply but, rather, that the widely suggested factors of limited light, nitrogen or phosphorus supply are the resource inputs which restrict productivity. Global change, by increasing atmospheric CO₂ partial pressure and global mean temperatures, is likely to increase the mean CO₂ concentration in the atmosphere, but the corresponding change in the oceans will be much less. There are, however, genotypic differences in the handling of inorganic C among the diversity of marine phytoplankton, and in impact on use of limiting nutrients, so increases in the mean CO₂ and HCO₃ ^- concentrations in surface ocean waters could cause changes in species composition. However, the rarity of inorganic C limitation of marine phytoplankton short-term photosynthesis, net productivity, or the maximum biomass, in today's ocean means that global change is unlikely to increase these three values in the ocean.     

太湖湖岸带浮游植物初级生产力特征及影响因素

利用高频溶解氧监测,估算了太湖梅梁湾湖岸带浮游植物初级生产力的高频变化特征。结合同步气象监测及浮游植物、浮游动物和营养盐的周年逐周观测数据,分析了气象和环境条件对富营养化湖泊浮游植物初级生产力的影响。结果发现,高频溶解氧监测估算的初级生产力变化与浮游藻类生物量的变化一致,能够反映出浮游植物生产力的昼夜变化、季节变化等规律。统计分析表明,气温对太湖这一富营养化湖泊初级生产力影响很大;氮的供给与浮游动物的选择性牧食也是影响浮游植物初级生产力的重要因素。湖岸带的水华堆积过程对初级生产力影响巨大,气象、水文过程又加剧了蓝藻水华初级生产力的变化幅度,反映出富营养化湖泊初级生产力可能存在极大的时空不均一性。研究表明,溶解氧高频监测法估算初级生产力能够捕捉到湖泊初级生产力的快速变化过程,可以用于富营养化湖泊初级生产力监测、蓝藻水华灾害预警中。     

The impact of phytoplankton on spectral water transparency in the Southern Ocean: implications for primary productivity

Spectral water transparency in the Northern Weddell Sea was studied during Austral spring. The depth of the 1-% surface irradiance level (euphotic depth) varied between 35 and 109 m and was strongly influenced by phytoplankton biomass. Secchi depths were non-linearly related to euphotic depth. In phytoplankton-poor water, the most penetrating spectral region was restricted to a relatively narrow waveband in the blue (488 nm), but the range was broader, between 488 and 525 nm when phytoplankton were abundant. Water transparency in the red spectral range was always low and only to a small extent affected by phytoplankton. Two independent procedures were used to quantify the impact of phytoplankton on spectral water transparency: (1) Regression analysis of spectral in situ vertical light attenuation coefficients in the sea, against coincident chlorophyll concentrations. This method gave chlorophyll-specific light attenuation coefficients; the y-intercept could be interpreted as a measure of light attenuation by pure water plus non-algal material. (2) Spectra of in vivo light absorption derived by spectroscopy, using phytoplankton enriched to varying degrees onto filters. Thus chlorophyll-specific absorption cross-sections were determined. Estimates obtained by both procedures were in close agreement. By integrating over the spectrum of underwater irradiance, in situ chlorophyll-specific absorption cross sections of phytoplankton suspensions, related to all photosynthetically active radiation, were calculated. Light absorption by phytoplankton for photosynthesis is accomplished mainly in the blue spectral range. Also dissolved and particulate organic matter contributed to the attenuation of blue light. Because in water poor in phytoplankton, underwater irradiance was progressively restricted to blue light, chlorophyll-specific absorption cross-sections of phytoplankton, averaged over the spectrum of photosynthetically active irradiance, increased with water depth. In water with elevated phytoplankton biomass, overall light attenuation was generally enhanced. However, because the spectral composition of underwater light changed relatively little with depth, except immediately below the water surface, light absorption cross-sections of phytoplankton changed little below 10 m depth. Vertical differences in the proportions of underwater light absorbed by the phytoplankton community here were mainly dependent on biomass variations. Because of the comparatively small attenuation of blue light by non-algal matter, the efficiency of light harvesting by phytoplankton at any given concentration of chlorophyll in Antractic waters is greater than in other marine regions. At the highest phytoplankton biomass observed by us, as much as 70% of underwater light was available for phytoplankton photosynthesis. When phytoplankton were scarce, <10% of underwater light was harvested by phytoplankton.Contribution within the European Polarstern Study (EPOS), supported by the Deutsche Forschungsgemeinschaft, Grant Ti 115/16-1 to MMT, the European Science Foundation, and by the Alfred Wegener Institut für Polar-und Meeresforschung, Bremerhaven     

太湖梅梁湾浮游植物叶绿素a和初级生产力

1998年5月～1999年8月对太湖梅梁湾4站点进行了每季1次、为期1年的初级生产力及相关因子研究,分析了梅梁湾叶绿素a含量和初级生产力的时空分布特征．结果表明,梅梁湾的叶绿素a含量、初级生产力均存在明显的季节变化和空间差异,春、夏季浮游植物叶绿索a含量和初级生产力要比秋、冬季高,空间上位于污染严重的直湖港口6#点叶绿索a含量和初级生产力要高,并大致呈现从湾内向湾口递减的趋势;在春、夏、秋季光照较强时,初级生产力最大指值出现在水下20～50cm处。到冬季垂直差异不明显;10～30℃之间初级生产力基本上随温度的上升而呈指数增加趋势．浮游植物生物现存量与初级生产力存在显著的正相关．营养盐与初级生产力相关性变化很大．光照显著地影响初级生产力的日变化。春、夏季强光作用下表面光抑制现象比较明显．     

Factors controlling phytoplankton primary productivity in Byram Lake,Mt. Kisco,N.Y., summer 1977

Phytoplankton productivity was measured in Byram Lake Reservoir during summer 1977. Depth integrated productivity (0–5 gC m^{– 2} d^–1) increased with station depth, which together with visibility measurements indicated that light did not limit deep station productivity (C1 and S2). Macrophytes at station C5 (shallow) reduced the euphotic zone to 0 in June.On a unit depth basis, C5 was the most productive station. Apparently changes in macrophyte growth, regulated by light and temperature, controlled phytoplankton production. At C1, productivity was related to levels of different nutrients at different depths, the thermocline influencing nutrient availability at mid-depth. At S2, NH₃-N controlled mid-depth productivity. Surface and mid-depth productivity appeared influenced by factors not measured in this study.     

Differential effects of changes in spectral irradiance on photoacclimation,primary productivity and growth in Rhodomonas salina (Cryptophyceae) and Skeletonema costatum (Bacillariophyceae) in simulated blackwater environments

The underwater light field in blackwater environments is strongly skewed toward the red end of the electromagnetic spectrum due to blue light absorption by colored dissolved organic matter (CDOM). Exposure of phytoplankton to full spectrum irradiance occurs only when cells are mixed up to the surface. We studied the potential effects of mixing‐induced changes in spectral irradiance on photoacclimation, primary productivity and growth in cultures of the cryptophyte Rhodomonas salina and the diatom Skeletonema costatum. We found that these taxa have very different photoacclimation strategies. While S. costatum showed classical complementary chromatic adaption, R. salina showed inverse chromatic adaptation, a strategy previously unknown in the cryptophytes. Transfer of R. salina to periodic full spectrum light (PFSL) significantly enhanced growth rate (μ) by 1.8 times and primary productivity from 0.88 to 1.35 mg C · (mg Chl^?1) · h^?1. Overall, R. salina was less dependent on PFSL than was S. costatum, showing higher μ and net primary productivity rates. In the high‐CDOM simulation, carbon metabolism of the diatom was impaired, leading to suppression of growth rate, short‐term ¹⁴C uptake and net primary production. Upon transfer to PFSL, μ of the diatom increased by up to 3‐fold and carbon fixation from 2.4 to 6.0 mg C · (mg Chl^?1) · h^?1. Thus, a lack of PFSL differentially impairs primarily CO₂‐fixation and/or carbon metabolism, which, in turn, may determine which phytoplankton dominate the community in blackwater habitats and may therefore influence the structure and function of these ecosystems.     

Effects of sediment resuspension on phytoplankton production: teasing apart the influences of light, nutrients and algal entrainment

1. Wind‐induced sediment resuspension can affect planktonic primary productivity by influencing light penetration and nutrient availability, and by contributing meroplankton (algae resuspended from the lake bed) to the water column. We established relationships between sediment resuspension, light and nutrient availability to phytoplankton in a shallow lake on four occasions. 2. The effects of additions of surficial sediments and nutrients on the productivity of phytoplankton communities were measured in 300 mL gas‐tight bottles attached to rotating plankton wheels and exposed to a light gradient, in 24 h incubations at in situ temperatures. 3. While sediment resuspension always increased primary productivity, resuspension released phytoplankton from nutrient limitation in only two of the four experiments because the amount of available nitrogen and phosphorus entrained from the sediments was small compared with typical baseline levels in the water column. In contrast, chlorophyll a entrainment was substantial compared with baseline water column concentrations and the contribution of meroplankton to primary production was important at times, especially when seasonal irradiance in the lake was high. 4. Comparison of the in situ light climate with the threshold of light‐limitation of the phytoplankton indicated that phytoplankton in the lake were only likely to be light‐limited at times of extreme turbidity (e.g. >200 nephelometric turbidity units), particularly when these occur in winter. Therefore, resuspension influenced phytoplankton production mainly via effects on available nutrients and by entraining algae. The importance of each of these varied in time. 5. The partitioning of primary productivity between the water column and sediments in shallow lakes greatly influences the outcome of resuspension events for water column primary productivity.     

Reconstruction of the Time Series of the Underwater Light Climate in a Shallow Turbid Lake

Lake ülemiste is a shallow, eutrophic lake which has served the city of Tallinn as a water reservoir for many centuries. Its light climate was studied by combining a routinely measured data set with a modelling approach. For 26 years (1978–2004), data was collected on such optically active substances (OAS) and water parameters as water colour, turbidity and phytoplankton biomass. Simple modelling enabled the quantification of long-term time-series data and the subsequent calculation of the diffuse attenuation coefficient, euphotic depth and average light of the mixed layer. Several changes in the hydrological cycle have taken place during the period under study, among which are an increase in the water level of about 0.5 m and a decrease in the external water load from 108 million m³ year⁻¹ to about 25 million m³ year⁻¹. At the same time euphotic depth has shown a distinct trend towards increasing since the early 1990s. The euphotic depth also showed an increase (from 1.1 to 1.4 m) due to an improvement in underwater light conditions – mainly in the spring (April and May) and autumn (October and November) because of the lower amount of dissolved organic matter in the lake. The average light availability in the mixed layer has increased, but this has not affected the phytoplankton biomass as the latter is not light-limited during the summer period.     

Photosynthèse phytoplanctonique: Réponse à un stimulus simple,imitant les variations rapides de la lumière engendrées par les vagues

Experiments were conducted in order to assess the response of marine phytoplankton production to a simplified stimulus modelling rapid light fluctuations induced by sea surface waves. Studies were carried out in the St Lawrence Estuary, during three clear and absolutely calm days. The photosynthesis of artificially incubated samples was generally depressed in fluctuating light, when compared with that in constant light of the same average intensity. There is, however, a daily trend of increased energy utilization in varying light, at saturating intensities, while some unpredictable instances of higher photosynthesis were observed in fluctuating light, especially at low intensities. Results are interpreted in terms of changes in carboxylation pathways, and might be a function of the average fluctuating light intensity and of the sun adaptation by the phytoplankton. It follows that the use of fixed light ‘imitated in situ’ incubators should be viewed with some caution.     

Influence of an Atypical Summer on the Primary Productivity of Phytoplankton in Two Tropical Ponds

Primary productivity of phytoplankton was measured in two fish ponds at monthly intervals during two annual cycles, of which the second one included an unusual drought. The temporal course of variations in phytoplankton primary productivity was essentially similar in both annual cycles, exhibiting peaks and troughs during summer and winter, respectively. However, monthly mean values of gross and net primary productivity of phytoplankton during the second year were several times higher than during the first. The greatest difference between the two years of investigation was found in the summer peak of primary productivity. It is estimated that an increase of nitrogen or phosphate or the N/P ratio by one unit during the second year caused a 2 to 16-fold enhancement of the rates of primary productivity. The seasonal changes of photosynthetic efficiency correlated with the concentrations of phosphate.     

DIEL VARIATIONS IN OPTICAL PROPERTIES OF MICROMONAS PUSILLA (PRASINOPHYCEAE)1

Micromonas pusilla (Butcher) Manton et Parke, a marine prasinophyte, was used to investigate how cell growth and division affect optical properties of phytoplankton over the light:dark cycle. Measurements were made of cell size and concentration, attenuation and absorption coefficients, flow cytometric forward and side light scattering and chl fluorescence, and chl and carbon content. The refractive index was derived from observations and Mie scattering theory. Diel variations occurred, with cells increasing in size, light scattering, and carbon content during daytime photosynthesis and decreasing during nighttime division. Cells averaged 1.6 μm in diameter and exhibited phased division, with 1.3 divisions per day. Scattering changes resulted primarily from changes in cell size and not refractive index; absorption changes were consistent with a negligible package effect. Measurements over the diel cycle suggest that in M. pusilla carbon‐specific attenuation varies with cell size, and this relationship appears to extend to other phytoplankton species. Because M. pusilla is one of the smallest eukaryotic phytoplankton and belongs to a common marine genus, these results will be useful for interpreting in situ light scattering variation. The relationship between forward light scattering (FLS) and volume over the diel cycle for M. pusilla was similar to that determined for a variety of phytoplankton species over a large size range. We propose a method to estimate cellular carbon content directly from FLS, which will improve our estimates of the contribution of different phytoplankton groups to productivity and total carbon content in the oceans.     

Environmental control of phytoplankton productivity in turbulent turbid systems

Factors affecting phytoplankton productivity are analysed in turbid systems, such as shallow lakes and rivers. When resuspension from the sediment or loading from the catchment significantly increases inorganic (non-algal) turbidity and hence light attenuation potentials for high production are not realised. Energy available for phytoplankton growth is strongly regulated by underwater light availability which depends on the critical mixing depth, fluctuating light intensities and algal circulation patterns. Higher production rates in shallow waters are often compensated by greater algal respiration due to higher water temperatures when compared to deeper lakes.Total daily integral production of turbulent, turbid environments can be predicted from a combination of easily measured variables such as maximum photosynthetic rates, algal biomass, surface irradiance and some measure of underwater light attenuation.