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.     

Vertical zonation and production rates of epiphytic algae on Phragmites australis

1. The vertical distribution of chlorophyll in epiphyton on Phragmites australis showed a peak in the middle sections of the submerged parts. Just below the water surface and above the sediment, chlorophyll concentrations were much less. 2. During winter and early spring, loosely attached diatoms were predominant just below the water surface and on the middle sections of the Phragmites plants. Near the bottom, adnate diatoms, parenchymatous thalli of chlorophytes and cyanobacteria were abundant. 3. At high photosynthetic active radiation (PAR) during May, filamentous species of Ulpthricophyceae and Zygophyceae developed dense populations on the middle sections of the stems. 4. Primary production rate was proportional to chlorophyll concentration although production maxima were recorded above the biomass maxima. 5. Where illumination was low, the chlorophyll-specific rate of photosynthesis (P^B rate) decreased proportionally with the vertical decrease of PAR in the littoral zone, independent of the chlorophyll concentration on the stems. 6. When illumination was high, the P^B rate decreased as biomass increased, and was independent of surface radiation. 7. The dependence of primary production rate on chlorophyll concentration produced a saturation curve with a maximum production at 4.6 μgC cm^?2h^?1.     

Growth and production ofPhragmites australis in Lake Vechten (the Netherlands)

Summary A study was carried out on several production characteristics ofPhragmites australis stands in Lake Vechten, the Netherlands. The aim was to quantify the role of this plant species in the lake's primary production and nutrient cycles (C, N and P).Maximum biomass values of 1247 and 286 g org.wt.m^–2 were measured in, respectively, shoots and underground organs in August. The shoot: root ratio was always>1. Plant litter accumulated particularly in spring and autumn. The relative growth rates of early shoots differed from those determined for the whole vegetation, indicating shoots of different ages in the latter. This was confirmed by counting the number of plants.The chemical composition of the plants changed with season. A carbon peak as high as 50.94% org.wt. was found in spring, which coincided with a nitrogen peak of 2.88% org. wt. At the end of the season N dissapeared faster from the shoots than C, whereas P declined from spring onwards.The production ofPhragmites contributed in 1979 for 5.8% to the lake's total primary production expressed in C, whereas the share of the total macrophytic unit amounted to 8.7%.The potential role ofPhragmites in the lake's nutrient cycles was largely due to nutrient leakage and decomposition.     

Contribution of photosynthetic sulphur bacteria to primary production in Lake Vechten

The role of photosynthetic sulphur bacteria as primary producers in monomictic Lake Vechten (The Netherlands) is described. Lake Vechten has a surface area of 4.7 ha, a maximum depth of 11.9 m and a mean depth of 6 m.Bacterial populations, appearing at the boundary layer of the oxidative and reductive zone from early June till late October, were composed of cyanobacteria, Chromatiaceae and green and brown coloured Chlorobiaceae. Predominating genera were Synechococcus, Chloronema, Chromatium and Thiopedia. The photosynthetic sulphur bacteria accounted for a primary production rate of 13.6–106.1 mg C.m^–2 day^–1, which corresponded to 3.9–17.5% of total daily productivity in the pelagial zone. The percentage of photosynthetic bacterial production of total annual planktonic primary production calculated for the entire pelagial zone, taking into account compensation for decreasing volume of lower strata, was 3.6% (i.e. 127 against a total production of 3 510 kg C.lake^–1yr^–1).     

Periphyton biomass,potential production and respiration in a shallow lake during winter and spring

Abundance, depth distribution, potential productivity and respiration of periphyton on short-time (1 month) and long-time incubated strips were followed monthly during the winter–spring (January–May) transition in a shallow eutrophic lake. A taxonomic shift occurred from dominance of diatoms under ice to chlorophyte dominance in spring communities on the long-time incubated strips, while diatoms dominated until May on the short-time incubated strips. Periphyton biomass accrual was low during the ice-covered winter months (November–January: 4 mg chl a m⁻² month⁻¹), but increased to a maximum of 112 mg chl a m⁻² month⁻¹ immediately after ice-out in February. During February–April, the biomass remained constant before declining in May. Periphyton on long-time incubated strips was equally distributed in the water column in winter (January–February), but was higher near the water surface in spring (March–May). Periphyton did not change with depth on the short-time incubated strips. The potential production to respiration ratio (P/R) was negatively correlated with periphyton biomass. Throughout the study, P/R was <1 for the short-time incubated periphyton, while this was only the case in March–April for the long-time incubations. This study showed a high productive capacity of winter periphyton, resulting in accumulation of a relatively high periphytic biomass early in the season. A massive periphyton density in eutrophic lakes already in winter–spring may potentially delay or prevent the establishment and re-occurrence of submerged macrophytes in the early oligotrophication phase following a reduction of the external nutrient loading. Handling editor: Luigi Naselli-Flores     

The aquatic macrophytes of Lake Vechten. Species composition,spatial distribution and production

The macrophytic species compositions in Lake Vechten of 1963 and 1979–80 were compared and showed a considerable change. The main vegetation types were mapped in 1973, 1978, 1979 and 1980. All macrophytic communities, i.e. submerged, floating-leaved and emergent vegetation types, declined mainly due to increasing water turbidity, increasing tree-shading and, from 1978 onwards, grazing and trampling by cattle.Production rates, derived from different combinations of measurement and calculation, were compared. The macrophytes contributed only about 7% to the total lake production in 1980.     

Seasonal changes in composition and biomass of epiphytic algae on the macrophyte Ranunculus penicillatus in a chalk stream,with estimates of production,and observations on the epiphytes of Cladophora glomerata

The epiphytic algae on surfaces of the macrophyte Ranunculus penicillatus (Dumort.) Bab. var. calcareus (R. W. Butcher) C. D. K. Cook and on Cladophora glomerata (L.) Kütz growing in the River Itchen at Otterbourne near Southampton were studied between February 1984 and June 1985. The river at this site has a mean flow rate of 0.33 m s^–1, and is about 16 m wide and on average 20 cm deep, with a discharge ranging through the year between 0.34 and 2.46 m³ s^–1. The pH of the river varies little around 8.2, with a mean alkalinity of 236 mg HCO _inf3 ^sup1 l^–1, because of its origin from chalk springs. Ranunculus grows throughout the year, with peaks of biomass in spring and autumn. It forms a very large surface for attachment of epiphytes, and covers on average 40% of the stream bed. Numbers of epiphyte cells removed from Ranunculus ranged through the year between 52 × 10³ and 271 × 10³ cells mm^–2 stream floor, with maximum numbers in April, and a secondary peak in October. This pattern partly reflects fluctuations in the biomass of Ranunculus; the number of cells per unit area of plant surface showed a broader spring peak and lower fluctuations in other seasons. Diatoms formed 65 to 98% of these epiphyte cells, with chlorophytes reaching their peak (10%) in summer and cyanophytes (25%) in autumn. Estimates of biomass of these epiphytic forms, derived from measurements of chlorophyll c, indicate a range between 30 and 100 g dry weight m^–2 of weed bed. Colonisation studies showed that the algae grow and reproduce throughout the year, with a mean generation time of about 5 days, suggesting an annual production of about 3 kg dry weight m^–2 of weed bed, which makes epiphytic algae the principal primary producers in the stream. The numbers and biomass of epiphytic algae on Cladophora are considerably less.The species of epiphytic algae found on Ranunculus were generally similar to those growing on Cladophora, and to epilithic algae on pebbles of the stream bed, but different species were dominant on the different substrata. Algal cells in the water column were all derived from benthic habitats, although their relative abundance was very different.     

Studies on Lake Erie's littoral algae; Host specificity and temporal periodicity of epiphytic diatoms

Substratum specificity and temporal periodicity of the attached diatom flora upon three aquatic vascular plants and an artificial substratum were examined in three Lake Erie marshes. No qualitative or quantitative specificity for substrata was observed. Variability of diatom assemblage structure within replicate samples of a particular substratum type was as great as, or greater than, variability between substrata. Diatom assemblages upon dowel rod displayed a mid to late summer density maximum. Variability of density maxima upon natural substrata was attributed to different growth rates of the host macrophytes. Diatom assemblages within each sampling site possessed a distinct temporal periodicity indicating that factors affecting diatom growth are heterogeneous in distribution throughout Lake Erie's littoral zone.     

Periphyton production in Fort River, Massachusetts

SUMMARY. The primary production and general ecology of a periphyton community of a New England, lowland stream were studied over a seventeen-month period. Temperature, light, periphyton chlorophyll-α, and community structure were monitored regularly. Seasonally distinct chlorophyll peaks coincided with the light maximum in early May, just prior to the appearance of leaves of riparian trees, and again in autumn after terrestrial leaf fall. During midwinter, despite low light and temperature levels and high stream discharge, mean chlorophyll concentrations remained similar to summer values.
A mathematical expression relating periphyton photosynthesis per unit chlorophyll-α to temperature, light and periphyton density was established with submersible light-dark chambers in situ . Survey data collected over the study period were employed in the empirical equation to estimate seasonal variations in periphyton primary production. Weekly mean daily estimates of periphyton gross production ranged from < 0.1 g O₂ m⁻², during midwinter, to 6.5 g O₂ m⁻² during early May. Estimated annual periphyton gross production and respiration were 0.58 and 1.27 kg O₂ m⁻², respectively. Factors influencing seasonal variations of Fort River periphyton standing crop are discussed.     

Diel patterns of phytoplankton productivity in Lake Vechten

Summary A review of the literature on phytoplankton primary production studies in lakes indicates that in most cases the daily primary production rates have been derived by multiplying the short-term (usually 3 or 4 hours) production measurement with a light factor. This factor is a ratio between the total irradiance during the day and during the period of incubation. The aim of the present study was to verify this method of calculation. Since the primary production rate is of great importance for the carbon cycle of lakes, this study can be regarded as a fundamental contribution to the integrated research of the ecosystem of lake Vechten (the Netherlands) (BESTet al., 1978).Photosynthetic carbon fixation was determined using the radiocarbon method of STEEMANN NIELSEN (1952). Thein situ measurements were carried out for 2 hours incubation periods starting at 15–30 minutes before sunrise and ending 15–30 minutes after sunset. Also measured were: water temperature, light penetration, oxygen and chlorophyll. Total incident solar radiation data were obtained from the Royal Netherlands Meteorological Institute (K.N.M.I.) at de Bilt (situated at 5 km from lake Vechten). The production during two consecutive 2-hourly periods,e.g. period 1 and period 2, were summed up and using the irradiance data during the 4 hours, daily production was calculated. These calculations were repeated for period 2 and 3, and so on. The sum of all the short-term measurements during a day,i.e. the total daily primary production, was taken for comparison with the production calculated by using the sum of the production during two successive periods and the light factor.Daily production rates obtained by extrapolating from short-term incubation periods both in the morning and in the late afternoon appeared to overestimate the real daily production (Fig. 1) up to 100%. It is likely, that in the morning the lack of nutrient limitation and relatively better physiological state of algae, and in the afternoon increase of phytoplankton biomass during the day can be considered as the factors causing the discrepancy between the calculated and real production. On the other hand, the underestimation of daily production rate based on midday time intervals on sunny days, was about 25%. It is attributed to photoinhibition, especially in the present study where the measurements were limited to the upper water strata (0–4m).Is is concluded that short-termin situ exposures between 10.00 and 14.00 hr appeared to be a good indicator of total primary production, although underestimates up to 25% on sunny days are to be taken into account.     

The effect of oxygen concentration on photosynthetic biomass production by algae

The effect of decreased oxygen concentration on photosynthetic biomass production was determined for Euglena gracilis Klebs strain z and Chlamydomonas reinhardtii Dangeard. At a constant carbon dioxide concentration of 0.03% (v/v), decreasing the oxygen concentration from 21% to 2% (v/v) gave a two-fold increase in dry-weight yield for E. gracilis; a result consistent with the operation of a functional glycollate pathway in this alga. A similar effect of oxygen concentration on dry-weight yield was not observed with C. reinhardtii.     

Instar-dependent mortality rates of coexisting Daphnia species in Lake Vechten, The Netherlands

In Lake Vechten, population parameters were determined for twocoexisting Daphnia species, D.hyalina and D.cucullata, fromApril 23 to December 10, 1986. Daphnia hyalina is dominant inspring and D.cucullata in summer and autumn. Size frequencydistributions were converted into instar frequency distributionsby using experimental growth curves and field measurements onnewborn size and the size at first reproduction. A discreteevent computer model was used to calculate the instar mortalityrates. Relative importances of instar mortality rates for thepopulation mortality rate were calculated and discussed. Themortality of the juvenile stages was found to be more importantthan adult mortality during most of the sampling season, despiteegg mortality when egg-bearing females are killed. Only in Septemberand October was the mortality of adult instars more important.Correlation coefficients between differences in the rates ofincrease and differences in mean brood size and instar mortalitiesshow that the former are caused mainly by differences in juvenilemortality, the main factor determining the replacement of D.hyalinaby D.cucullata during early summer.     

Empirical and dynamical models of production and biomass of benthic algae in lakes

This work presents new empirical and dynamical models for benthic algae in lakes. The models were developed within the framework of a more comprehensive lake ecosystem model, LakeWeb, which also accounts for phytoplankton, bacterioplankton, two types of zooplankton (herbivorous and predatory), macrophytes, prey fish and predatory fish. The new dynamic model provides seasonal variations (the calculation time is 1 week). It is meant to account for all factors regulating the production and biomass of benthic algae for lakes in general. This work also presents and uses a new data-base established by us from published sources. Many of the lakes included in this study are situated in the former Soviet Union. They were investigated during the Soviet period and the data and results have up until now been largely unknown in the West. We present empirical models for benthic algae, and show that the biomass of benthic algae in whole lakes can be estimated from the ratio between the lake area above the Secchi depth to the total lake area and the primary production of phytoplankton. We also present several critical tests of the dynamical model. The dynamical and empirical models give corresponding results over a wide limnological domain. We provide algorithms for (1) the production rate of benthic algae (2) the elimination rate (related to the turnover time of benthic algae), (3) the rate of benthic algae consumption by zoobenthos, and (4) the rate of physical erosion of benthic algae. Our results indicate that the production of benthic algae is highly dependent on lake morphometry and sediment character, as well as water clarity, and less dependent on nutrient (phosphorus) concentrations in water and sediments. This work provides new quantitative support to such conclusions and also a useful model for predictions of production and biomass of benthic algae.     

The diversity,biomass and production of zooplankton in Lake Inarijärvi

About 650 zooplankton samples were collected from Lake Inarijärvi in 1977–1979 from the littoral and pelagial zones of the lake. One hundred and twenty-three zooplankton taxa were found and most of them can be considered euplanktonic.The most important species were Holopedium gibberum, Daphnia cristata, Cyclops spp. and Eudiaptomus spp. Mean pelagial zooplankton biomass was 0.29 g m^–3 in the 0–5 m depth zone, 0.17 g m^–3 in 5–10 m and 0.11 g m^–3 in 10–20 m.The zooplankton biomass at a sandy shore was about 0.09 g m^–3, at a stony shore 0.05 g m^–3 and at a vegetated shore 0.76 g m^–3. About 70% of the whole zooplankton production consisted of crustaceans.The sum of herbivore and carnivore zooplankton production in the pelagial area during the summer was 210–330 kg ha^–1 × 3 months.     

Spatial and temporal variation in the biomass and nutrient status of epilithic algae in Lake Erken, Sweden

1. The aim of this study was to estimate patchiness in biomass and in the internal nutrient status of benthic algae on hard substrata (epilithon) in Lake Erken, Sweden, over different levels of distance, depth and time. Knowledge of the sources and scale of patchiness should enable more precise estimation of epilithic biomass and nutrient status for the entire lake. We focused on the horizontal scale, about which little is known. 2. We sampled epilithon by SCUBA diving and used a hierarchical sampling design with different horizontal scales (cm, dm, 10 m, km) which were nested in two temporal scales (within and between seasons). We also compared two successive years and three sampling depths (0, 1 and 4 m). Biomass was measured as particulate carbon and chlorophyll a (Chl a) and internal nutrient status as carbon : nitrogen : phosphorus (C : N : P) ratios and as specific alkaline phosphatase activity (APA). 3. Horizontal variation accounted for 60–80 and 7–70% of the total variation in biomass and in nutrient status, respectively, at all depths and during both years. Both small and large scales accounted for significant variation. We also found variation with time and depth. Biomass increased in autumn after a summer minimum, and the within‐season variation was very high. The lowest biomass was found at 0 m depth. Both N and P limitation occurred, being higher in 1996 than in 1997 and decreased with depth. 4. As a consequence, any sampling design must address variation with distance, depth and time when estimating biomass or nutrient limitation of benthic algae for an entire lake. Based on this analysis, we calculated an optimal sampling design for detecting change in the epilithic biomass of Lake Erken between different sampling days. It is important to repeat the sampling as often as possible, but also the large scales (10 m and km) and the dm scale should be replicated. Using our calculations as an example, and after a pilot study, an optimal sampling design can be computed for various objectives and for any lake. 5. Short‐term impact of the wind, light and nutrient limitation, and grazing, might be important in regulating the biomass and nutrient status of epilithic algae in Lake Erken. Patchiness in the nutrient status of algae was not coupled to the patchiness of biomass, indicating that internal nutrients and biomass were regulated by different factors.     

The bacterioplankton of Lake Taihu,China: abundance,biomass, and production

Abundance, biomass and production of pelagic bacteria were examined over one year at monthly sampling intervals across a trophic profile in Meiliang Bay, Lake Taihu. With the lowest density in the open lake, the bacterial abundance showed a clear trend in relation to trophic status. The carbon content per cell was higher in autumn and winter, and the opposite was true for bacterial biomass. Bacterial ³[H]-TdR and ¹⁴[C]-Leu incorporation rates, cell production, turnover times and carbon production varied during the annual cycle at different sites. The ratio of bacterial production to primary production was high, independently of the method used, indicates that the microbial food web in Lake Taihu is an important component of the total food web of the lake and dominated by external inputs. Electronic supplementary material Supplementary material is available for this article at and is accessible for authorized users     

Carbon flow across the sediment-water interface in Lake Vechten,The Netherlands

The turnover and exchange rates, as well as the diffusion processes, concerning the input and output of carbon compounds at the mud-water interface, were studied. The carbon input rates were derived from the annual sedimentation rates of particulate organic matter (about 1 100 kg C · yr⁻¹). The nature of the sedimented POC, and its breakdown pathways and turnover rates towards important metabolic intermediates in methanogenesis, were examined. The breakdown kinetics ofChlorella cell walls, a dominant green alga in Lake Vechten, was studied using U-¹⁴C-labelled cell walls. The breakdown of the cell walls appears to the rate-limiting step in anaerobic mineralization. Using first order kinetic equations, and HPLC and GLC and radio-chemical methods, turnover rate constants (k-values) of between 0.18 and 0.32 day⁻¹ and pool sizes of algal cell walls of 37 to 80 μg · g⁻¹ wet mud were found, giving turnover rates of 7.7 to 25.6 μg · g⁻¹ · day⁻¹ of cell wall material. The turnover rates (k-values between 0.07 and 0.31 h⁻¹) of acetate, the most important breakdown product, and its concentration gradients (between 5 and 30 μmol) and diffusion coefficient (D_s = 2.2 × 10⁻⁶ cm² · s⁻¹) just in and above mud-water interface, was quantified. The diffusion of acetate, within the sediments, could not account for the turnover rates observed. Finally, from acetate flux data and from those on the rates of formation of carbon dioxide and methane, the output of carbon and its cycling in Lake Vechten are discussed.     

海草附生藻类生物量的主要影响因子

海草叶片上的附生藻类负荷量很大程度上决定了其对海草影响的强度及利弊的方向,它是各种非生物因子及生物因子相互作用的结果.总结了近20a来海草附生藻类生物量控制因子的研究进展,归纳得出影响海草附生藻类生物量的因子主要有生物因子中海草自身的生理特性和食草动物的摄食作用、化学因子中的营养盐浓度、物理因子中的可利用光强度和水流等.其中,作为主要上行效应的营养盐浓度以及作为主要下行效应的摄食作用对海草附生藻类生物量起十分重要的调节作用.最后,对未来的研究提出了展望:①上行效应及下行效应对于调节海草-附生藻类群落结构的相对重要性及其定量分析;②在各种因子单独作用以及联合作用下附生藻类生物量的变化规律;③海草释放化感物质调节附生藻类生长的机制;④附生藻类对海草生长的综合作用机制.