Linking elements to biochemicals: effects of nutrient supply ratios and growth rates on fatty acid composition of phytoplankton species

Three species of marine phytoplankton, Rhodomonas sp., Isochrysis galbana Parke, and Phaeodactylum tricornutum Bohlin, were cultivated in semicontinuous cultures to test biochemical responses (fatty acids; FAs) to five nitrogen (N):phosphorus (P) supply ratios and four growth rates (dilution rates). The characteristic FA profile was observed for each algal species (representing particular algal class), which remained relatively stable across the entire ranges of N:P supply ratios and growth rates. For all species, significant direct effects of N:P supply ratios on FAs were found at lower growth rates. The highest saturated and monounsaturated fatty acid (SFA and MUFA) contents were observed under N deficiency at the lowest growth rate in all three species, while responses of polyunsaturated fatty acids (PUFAs) revealed no consistent pattern. Total FAs (and SFAs and MUFAs) in all species showed significant negative correlations with N cell quota (Q_N) under N deficiency, but PUFAs had species‐specific correlations with Q_N. The results show that characteristic FA profiles of algal genus or species (representing particular algal classes) underlie fluctuations according to culture conditions. The significant correlation between FAs and Q_N under N deficiency suggests that elemental and biochemical limitation of phytoplankton should be considered mutually as determinants of food quality for zooplankton in marine ecosystems.     

INTENSE GRAZING AND PREY‐DEPENDENT GROWTH OF PFIESTERIA PISCICIDA (DINOPHYCEAE)1

Grazing and growth of Pfiesteria piscicida (Pfiest) were investigated using batch and cyclostat cultures with Rhodomonas sp. (Rhod) as prey. Observed maximum growth rates (1.4 d^?1) and population densities (2 × 10⁵ cells·mL^?1) corresponded to values predicted by Monod functions (1.76 d^?1; 1.4 × 10⁵ cells·mL^?1). In batch cultures under a range of prey‐to‐predator ratios (0.1:1 to 180:1) and prey concentrations (1000–71,000 cells·mL^?1), Rhodomonas sp. was always depleted rapidly and P. piscicida concentrations increased briefly. The rate of Rhodomonas sp. depletion and the magnitude of P. piscicida population maxima depended on the prey‐to‐predator ratio and prey concentration. Starvation resulted in cell cycle arrest at G1 and G2+M and ultimately the demise of both P. piscicida and Rhodomonas sp. populations, demonstrating the dependence of P. piscicida on the supply of appropriate prey. The depletion of Rhodomonas sp. populations could be attributed directly to grazing, because P. piscicida did not exert detectable inhibitory effects on the growth of Rhodomonas sp. but grazed intensely, with maximum grazing rates>10 Rhod·Pfiest^?1·d^?1 and with no apparent threshold prey abundance for grazing. The results suggest that 1) the abundance of appropriate prey may be an important factor regulating P. piscicida abundance in nature, 2) P. piscicida may control prey population, and 3) high growth and grazing potentials of P. piscicida along with cell cycle arrest may confer survival advantages.     

GROWTH RATE VARIATION IN THE N:P REQUIREMENT RATIO OF PHYTOPLANKTON1

Theoretical considerations predict that the cell N:P ratio at transition from nitrogen limitation to phosphorus limitation of phytoplankton growth (critical ratio, R_c) varies, as a function of population growth rate. This prediction is confirmed by experimental, data from the literature along with new experimental data for the marine, prymnesiophyte Pavlova lutheri (Droop) Green. R_c passes through a maximum at intermediate growth rates for the three phytoplankton species for which data, are available, but there is significant interspecific variability in its value. There is no theoretical or experimental evidence to support the idea that the ratio of subsistence N and P cell quotas is equal to R_c over the range of growth rates, or that the subsistence quota ratio equals the ratio of the N and P cell quotas minus a storage fraction. Examination of N:P composition ratios can be used to determine which nutrient is limiting, but cannot be used to determine relative growth rates or competitive advantage between species limited by the same nutrient. Growth rates are determined by environmental conditions and by the cell quota of the limiting nutrient, without reference to the cell quota of the non-limiting nutrient.     

Ingestion rates and dietary lipids affect growth and fatty acid composition of Dendraster excentricus larvae

This study investigated the effect of single and mixed algal diets on growth, fatty acid composition and ingestion rates for Dendraster excentricus larvae. Larvae were assigned to three single algal diet treatments Isochrysis galbana, Dunaliella tertiolecta or Rhodomonas sp. and four mixed algal diet treatments D. tertiolecta and Rhodomonas, I. galbana and D. tertiolecta, I. galbana and Rhodomonas, D. tertiolecta, Isochrysis galbana and Rhodomonas sp. Small amounts (0.36-0.6%) of stearic acid (18:0) were seen in the three algae used but a relatively large percentage (7-25%) of this SAFA was found in Dendraster larvae. The alga D. tertiolecta had the highest percentage (51.7%) of the short chain polyunsaturated fatty acid (PUFA) linolenic acid 18:3(n-3) and trace amounts (0.02-0.14%) of the long chain PUFAs eicosapentanoic (EPA, 20:5(n-3) and docosahexanoic acids (DHA, 22:6(n-3)). However, sand dollar larvae demonstrated the ability to elongate and desaturate shorter chain (18 carbon) polyunsaturated fatty acids (PUFA) to longer chain (20 carbon) n-3 PUFA. Thus high levels of 18:3(n-3) in D. tertiolecta led to high levels of EPA and low levels of 18:3(n-3) in Dendraster larvae fed this diet. Rhodomonas sp. had the highest percentage of stearidonic acid (18:4(n-3), 38.14%) and EPA (10.6%). Despite high levels of 18:4(n-3) in Rhodomonas sp. this acid was absent or found at very low levels in larvae fed this alga, or any combination of this alga. I. galbana had the highest percentage of DHA(14.3%) but was almost devoid of EPA (0.43%). Although Rhodomonas sp. and I. galbana had high levels of EPA and DHA sand dollar larvae did not incorporate higher levels of these long chain PUFAs into their lipids compared to those fed the alga D. tertiolecta. Dendraster larvae synthesized a number of 20 and 22 carbon non-methylene interrupted dienes (NMID), with levels increasing with larval stage. Higher ingestion rates were observed for Dendraster larvae fed single algal diets (Rhodomonas sp. or D. tertiolecta) and lower ingestion rates for those fed mixed algal diets. The highest ingestion rates were for 8-arm Dendraster larvae fed the large alga Rhodomonas sp. presented as a single algal diet. When fed a combination of three algae, selection of particles varied slightly depending on stage with 8-arm larvae ingesting slightly more of the larger algal cell in the mixture than 6-arm larvae. The present study suggests that regardless of the ratios given larvae might have an optimum ratio of different sized particles at which they can feed. The mixed algal diet of I. galbana and D. tertiolecta was the best algal diet leading to significantly larger larvae with high survival and development to metamorphosis. The single algal diets of Rhodomonas sp. or Dunaliella tertiolecta were the second and third best algal diets based on growth and survival to metamorphosis.     

PHOSPHATE‐LIMITED GROWTH OF PAVLOVA LUTHERI (PRYMNESIOPHYCEAE) IN CONTINUOUS CULTURE: DETERMINATION OF GROWTH‐RATE‐LIMITING SUBSTRATE CONCENTRATIONS WITH A SENSITIVE BIOASSAY PROCEDURE1

The relationship between steady‐state growth rate and phosphate concentration was studied for the marine prymnesiophyte Pavlova lutheri (Droop) J. C. Green grown in a chemostat at 22°C under continuous irradiance. A bioassay procedure involving short‐term uptake of 10 picomolar spikes of ³³P‐labeled phosphate was used to estimate the concentration of phosphate in the growth chamber. The relationship between growth rate and phosphate was well described by a simple rectangular hyperbola with a half‐saturation constant of 2.6 nM. The cells were able to take up micromolar spikes of phosphate at rates two to three orders of magnitude higher than steady‐state uptake rates. The kinetics of short‐term uptake displayed Holling type III behavior, suggesting that P. lutheri may have multiple uptake systems with different half‐saturation constants. Chl a:C ratios were linearly related to growth rate and similar to values previously reported for P. lutheri under nitrate‐limited conditions. C:N ratios, also linearly related to growth rate, were consistently lower than values reported for P. lutheri under nitrate‐limited conditions, a result presumably reflecting luxury assimilation of nitrogen under phosphate‐limited conditions. C:P ratios were linearly related to growth rate in a manner consistent with the Droop equation for growth rate versus cellular P:C ratio.     

Effects of zooplankton on nutrient availability and seston C : N : P stoichiometry in inshore waters of Lake Biwa, Japan

Forty-eight-hour experimental manipulations of zooplankton biomass were performed to examine the potential effects of zooplankton on nutrient availability and phytoplankton biomass (as measured by seston concentration) and C : N : P stoichiometry in eutrophic nearshore waters of Lake Biwa, Japan. Increasing zooplankton, both mixed-species communities and Daphnia alone, consistently reduced seston concentration, indicating that nearshore phytoplankton were generally edible. The zooplankton clearance rates of inshore phytoplankton were similar to rates measured previously for offshore phytoplankton. Increased zooplankton biomass led to increased concentrations of nutrients (NH₄-N, soluble reactive phosphorus [SRP]). Net release rates were higher than those found in previous measurements made offshore, reflecting the nutrient-rich nature of inshore seston. Zooplankton nutrient recycling consistently decreased TIN : SRP ratios (TIN = NH₄ + NO₃ + NO₂). This effect probably resulted from the low N : P ratios of nearshore seston, which were lower than those commonly found in crustacean zooplankton and thus resulted in low retention efficiency of P (relative to N) by the zooplankton. Thus, zooplankton grazing inshore may ameliorate algal blooms due to direct consumption but tends to create nutrient supply conditions with low N : P, potentially favoring cyanobacteria. In comparison with previous findings for offshore, it appears that potential zooplankton effects on phytoplankton and nutrient dynamics differ qualitatively in inshore and offshore regions of Lake Biwa. Received: September 4, 2000 / Accepted: January 23, 2001     

STEADY STATE GROWTH OF PHYTOPLANKTON IN CONTINUOUS CULTURE: COMPARISON OF INTERNAL AND EXTERNAL NUTRIENT EQUATIONS1,2

Previously, there have been conflicts over whether external or internal nutrient concentrations control phytoplankton growth rates at steady state in continuous culture. To experimentally demonstrate that both equations equally describe steady state growth, continuous culture studies with phosphorus-limited growth of the chrysophyte Monochrysis lutheri Droop were carried out over the entire growth rate region up to biomass washout. Data were examined using both the Monod and Droop equations, and, even though there were significant variations in the yield coefficient with growth rate, the data fit both equations reasonably well. Because of their relative simplicity, the Droop equation and an equation combining both the Monod and Droop equations are better suited for expressing kinetic data than the Monod equation. It is crucial, though, that the criteria necessary to achieve steady state be fulfilled.     

EFFECTS OF VARIATION IN TEMPERATURE. I. ON THE BIOCHEMICAL COMPOSITION OF EIGHT SPECIES OF MARINE PHYTOPLANKTON1

The influence of temperature on the biochemical composition of eight species of marine phytoplankton was investigated. Thalassiosira pseudonana Hasle and Heim-dal, Phaeodactylum tricornutum Bohlin and, Pavlova lutheri Droop (three of eight species studied) had minimum values of carbon and nitrogen quotas at intermediate temperatures resulting in a broad U-shaped response in quotas over the temperature range of 10 to 25°C. Protein per cell also had minimum values at intermediate temperatures for six species. For T. pseudonana, P. tricornutum, and P. lutheri, patterns of variation in carbon, nitrogen, and protein quotas as a function of temperature were similar. Over all species, lipid and carbohydrate per cell showed no consistent trends with temperature. Only chlorophyll a quotas and the carbon: chlorophyll a ratios (θ) showed consistent trends across all species. Chlorophyll a quotas were always lower at 10°C than at 25°C. Carbon: chlorophyll a ratios (θ) were always higher at 10°C than at 25°C. We suggest that although θ consistently increases at lower temperatures, the relationship between temperature and θ ranges from linear to exponential and is species specific. Accordingly, the interspecific variance in θ that results from species showing a range of possible responses to temperature increases as temperature declines and reaches a maximum at low temperatures. High photon flux densities appear to increase the potential interspecific variance in the carbon: chlorophyll a ratio and therefore exacerbate these trends.     

STEADY-STATE LUXURY CONSUMPTION AND THE CONCEPT OF OPTIMUM NUTRIENT RATIOS: A STUDY WITH PHOSPHATE AND NITRATE LIMITED SELENASTRUM MINUTUM (CHLOROPHYTA)1

Selenastrum minutum (Naeg.) Collins was grown over a wide range of growth rates under phosphate or nitrate limitation with non-limiting nutrients added to great excess. This resulted in saturated luxury consumption. The relationships between growth rate and cell quota for the limiting nutrients were well described by the Droop relationship. The observed variability in N cell quota under N limitation as reflected in k_Q·Q_max^?1*, was similar in magnitude to previously reported values but k_Q·Q_max^?1* for P under P limitation was greater than previously reported for other species. These results were evaluated in light of the optimum ratio hypothesis. Our findings support previous work suggesting that the use of a single optimum ratio (k_Qi·K_Qj^?1) is inappropriate for dealing with a species growing under steady-state nutrient limitation. Under these conditions the optimum ratio should be viewed as a growth rate dependent variable. Two approaches for testing the growth rate dependency of optimum ratios are proposed. The capacity for luxury consumption differed between nutrients and was growth rate dependent. At low growth rates, the coefficient of luxury consumption (R_sat) for P was ca. four times that for N. The set of all possible relationships between N and P cell quota under these conditions was reported and these values were then used to establish the cellular N:P niche boundaries for S. minutum. Cell quotas of non-limiting nutrients were not described by the Droop equation. Analysis showed that as the cellular N:P ratio deviates from the optimum ratio, the ability of the Droop equation to describe the relationship between growth rate and non-limiting cell quotas decreases. When non-limiting nutrient cell quotas are saturated, the Droop equation appears to be invalid. Previously reported patterns of non-limiting nutrient utilization are summarized in support of this conclusion. The physiological and ecological consequences of luxury consumption and growth rate dependent optimum ratios are considered.     

NITRATE AND PHOSPHATE UPTAKE INTERACTIONS IN A MARINE PRYMNESIOPHYTE1

The short- and long-term uptake of nitrate and phosphate ions, and their interactions, were studied as functions of the preconditioning of Pavlova lutheri (Droop) Green. Populations were preconditioned in continuous culture at a variety of growth rates and N:P supply ratios. The maximum uptake rates cell^?1 for nitrate and phosphate were of similar magnitudes, in spite of the forty-fold smaller requirement for phosphorus. Short-term phosphate uptake was independent of the nitrate concentration, but the short-term nitrate uptake rate was reduced in the presence of phosphate. The severity of inhibition of nitrate uptake by phosphate was positively correlated with the preconditioning N:P supply ratio and the preconditioning growth rate. In response to large additions of nutrients, P. lutheri was able to increase its phosphorus content sixty-fold, but was only able to take up enough nitrate to double its nitrogen content. The high rate of phosphate uptake relative to its requirement, the inhibition of nitrate uptake by phosphate, and the large capacity for phosphorus storage relative to its requirement, all of which were observed even under N limitation, may imply that even where nitrogen is limiting there can be interspecific competition for available phosphate.     

Effects of nutrient recycling by zooplankton and fish on phytoplankton communities

In laboratory experiments we tested the hypothesis that nutrients supplied by fish and zooplankton affect the structure and dynamics of phytoplankton communities. As expected from their body size differences, fish released nutrients at lower mass-specific rates than Daphnia. On average, these consumers released nutrients at similar N:P ratios, although the ratios released by Daphnia were more variable than those released by fish. Nutrient supply by both fish and Daphnia reduced species richness and diversity of phytoplankton communities and increased algal biomass and dominance. However, nutrient recycling by fish supported a more diverse phytoplankton community than nutrient recycling by Daphnia. We conclude that nutrient recycling by zooplankton and fish have different effects on phytoplankton community structure due to differences in the quality of nutrients released. Received: 21 December 1998 / Accepted: 31 May 1999     

Food quality for Daphnia in humic and clear water lakes

1. Growth and reproduction of Daphnia fed lake seston were measured in two categories of meso‐ to eutrophic lakes differing with respect to terrestrial organic matter influence (humic and clear water lakes). The content of highly unsaturated fatty acids (HUFA), P and N, as well as the taxonomical composition of seston were analysed. 2. Seston HUFA and C : P ratios were similar between lake categories, whereas C : N ratios were lower in the clear water lakes in both spring and summer. Despite the similarity in HUFA and P content of seston, Daphnia growth rate, clutch size and the proportion of gravid females were, respectively, about 1.5, 3 and 6 times higher in the clear water lakes. 3. Differences in growth and reproduction were related to a combination of higher N content and good fatty acid quality of the seston in the clear water lakes. Relatively high biomass of edible algae, such as Rhodomonas sp. and Cryptomonas sp., in the clear water lakes, and differences in water pH likely contributed to the observed differences in Daphnia growth and reproduction between lake categories. Additionally, it is possible that Daphnia was energy limited in the humic lakes despite high particulate organic carbon (POC) concentrations, as the contribution of non‐algal and detrital C to the POC pool was high. 4. Our results suggest that dietary HUFA content has the potential to improve herbivore growth and reproduction if N and P are not limiting. N merits more attention in studies of zooplankton nutrition.     

EFFECT OF NUTRIENT LIMITATION ON FATTY ACID AND LIPID CONTENT OF MARINE MICROALGAE1

Phaeodactylum tricornutum and Chaetoceros sp. (Badllariophyceae), Isochrysis galbana (clone T-Iso) and Pavlova lutheri (Prymnesiophyceae), Nannochloris atomus (Chlorophyceae), Tetraselmis sp. (Prasinophyceae), and Gymnodinum sp. (Dinophyceae) were cultured at different extents of nutrient-limited growth: 50 and 5% of μ_max. The lipid content of the algae was in the range 8.3–29.5% of dry matter and was generally higher in the Prymnesiophyceae than in the Prasinophyceae and the Chlorophyceae. Increasing extent of phosphorus limitation resulted in increased lipid content in the Bacillariophyceae and Prymnesiophyceae and decreased lipid content in the green flagellates N. atomus and Tetraselmis sp. The fatty acid composition of the algae showed taxonomic conformity, especially for the Bacillariophyceae, where the major fatty adds were 14:0, 16:0, 16:1, and 20:5n-3. These fatty acids were dominant also in the Prymnesiophyceae together with 22:6n-3. An exception was I. galbana, in which 18:1 was the major monounsaturated fatty add and 20:5n-3 was absent. The fatty acids of N. atomus and Tetraselmis sp. varied somewhat, but 16:0, 16:1, 18:1, 18:3n-3, and 20:5n-3 were most abundant. Gymnodinum sp. contained mainly 16:0, 18:4n-3, 20: 5n-3, and 22:6n-3. An increased level of nutrient limitation (probably phosphorus) resulted in a higher relative content of 16:0 and 18:1 and a lower relative content of 18:4n-3, 20:5n-3, and 22:6n-3. The nutrient limitation probably reduced the synthesis of n-3 polyunsaturated fatty acids.     

CHRONIC EFFECTS OF ULTRAVIOLET-B RADIATION ON GROWTH AND CELL VOLUME OF PHAEODACTYLUM TRICORNUTUM (BACILLARIOPHYCEAE)1

Cultures of the temperate estuarine diatom, Phaeodactylum tricornutum Bohlin (NEPCC Clone 31), were grown under ambient intensities of ultraviolet-A radiation (UVAR), photosynthetically active radiation (PAR), and various intensities of ultraviolet-B radiation (UVBR; 290–320 nm). Growth rates and cell volumes were monitored for 36 d. UVBR decreased growth rates and increased cell volumes. Sensitivity of growth to UVBR increased with time. Growth rates of P. tricornutum decreased with increasing ratios of UVBR:UVAR + PAR.     

Oil production from five marine microalgae for the production of biodiesel

Marine microalgae were studied as potential resources for the production of biodiesel. Five marine microalgae, Tetraselmis suecica, Phaeodactylum tricornutum, Chaetoceros calcitrans, Isochrysis galbana, and Nannochloropsis oculata were cultured in f/2 media, 12:12 L:D cycle at 20 ± 1°C with a light intensity of 36.3 μmol/m²/sec using a 15-L circular cylindrical photobioreactor. The dry cell weight, specific growth rate, biomass productivity, oil content and fatty acid composition of palmitic acid, stearic acid, oleic acid, linoleic acid, and linolenic acid of microalgae were determined. T. suecica, I. galbana, and N. oculata showed high dry cell weights of 0.58, 0.57, and 0.57 g/L, respectively. The culture period of T. suecica to reach the stationary phase was 9 days. On the other hand, N. oculata showed the longest culture period of 28 days to reach the stationary phase. T. suecica absorbed nitrate at the initial stages of cell growth, decreasing the nitrate concentration to 0.5 mg/L on day-7 of the culture. The highest oil contents were observed in P. tricornutum with a 25.31% dry cell weight and I. galbana with a 23.15% dry cell weight on day-9 after the stationary phase. I. galbana showed 417.33 mg of palmitic acid per g oil and T. suecica showed 235.61 mg of oleic acid per g oil. Stearic acid, linoleic acid, and linolenic acid did not exceed 30.02 mg/g oil in any of the microalgae. T. suecica showed the shortest culture period of 9 days to reach the stationary phase. Therefore, the highest biomass production of 0.58 g/L was obtained and I. galbana showed high biomass production of 0.57 g/ L and oil content of 23.15% of dry cell weight. Therefore, T. suecica and I. galbana can be selected as a potential candidate for the production of biodiesel.     

EFFECTS OF FLOWING WATER ON NITROGEN- AND PHOSPHORUS-LIMITED PHOTOSYNTHESIS AND OPTIMUM N:P RATIOS BY SPIROGYRA FLUVIATILIS (CHAROPHYCEAE)1,2

The effects of flowing water on net photosynthesis, dark respiration, specific growth rate, and optimum N:P ratios by Spirogyra fluviatilis Hilse were assessed. The alga was cultivated under nitrogen or phosphorus limitation in laboratory streams at three flow velocities: 3, 12, and 30 cm·s^?1. The Droop equation adequately described respiration and photosynthesis (PS_net) as a function of N or P cell quota (Q^N or Q^p). The data show that for N- or P-limited Spirogyra fluviatilis, flowing water is physiologically costly. Generally, flowing water had little effect on respiration rates; however, the proportion of gross photosynthesis devoted to dark respiration did increase with flow velocity. For photosynthesis, the minimum N and P cell quotas increased with velocity, and the theoretical PS_net maxima for N and P both appeared greatest at 12 cm·s^?1. The Droop models showed that for any given Q^N or Q^p, PS_net, was reduced by the 30-cm·s^?1 treatment. Consistent with this finding, independent estimates of specific growth rates for P-limited S. fluviatilis in the laboratory streams were inversely related to flow velocity when ambient PO₄^?3 was undetectable. However, growth was not diminished at the fastest velocity when PO₄^?3 was available for uptake. Thus, the increase in cellular phosphorus demand can be offset by flow-enhanced P uptake when conditions permit; otherwise, growth will be impaired. The optimum N:P ratios for S. fluviatilis at 3, 12, and 30 cm·s^?1 were 50, 58, and 52 by atoms, respectively, when calculated for PS_net= 0. The optimum ratios were inversely related to PS_net and decreased to approximately 20 when PS_net was near maximum. The potential for flowing water to mediate nutrient partitioning among lotic algae by altering growth rates and optimum nutrient ratios is discussed.     

Grazing and growth responses of a marine oligotrichous ciliate fed with two nanoplankton: does food quality matter for micrograzers?

To investigate whether predator growth and grazing would depend on prey properties besides size, we studied the numerical and functional responses of a marine oligotrichous ciliate isolated in Hong Kong coastal waters, Strobilidium sp., on two nanoplanktonic preys of similar size. The growth and ingestion rates of Strobilidium sp. could be fit with prey concentrations by hyperbolic curves. Strobilidium sp. exhibited higher maximal growth rates and gross growth efficiencies, and lower maximal clearance rates on Nannochloropsis sp. than on Isochrysis galbana. Our results demonstrate that prey properties presumably food quality can have a considerable effect on predator growth and grazing and implications on phytoplankton community structure and biogeochemical cycling.     

Phytoplankton community responses to nutrient and iron enrichment under different nitrogen to phosphorus ratios in the northern Baltic Sea

The impact of nutrient enrichment on the phytoplankton community structure, and particularly cyanobacteria, was studied in a 3-week mesocosm experiment conducted in August 2001 in the Archipelago Sea, a part of the northern Baltic Sea. The factorial design experiment included daily additions of nitrogen (N) and phosphorus (P) at two mass ratios, 1N:1P and 7N:1P, respectively, additions of iron (Fe) and a synthetic chelator, ethylenediaminetetraacetic acid (EDTA). The floating enclosures (400 l) were sampled for analyses of phytoplankton biomass and community structure, phytoplankton primary production, chlorophyll a, nutrients, and hepatotoxins. Chlorophyll a concentration, phytoplankton biomass and primary production increased most in the 7N:1P treatment. The increase was mainly due to an abundant growth of chlorophytes (Dictyosphaerium subsolitarium, Kirchneriella spp., Monoraphidium contortum, and Oocystis spp.), pennate diatoms (especially Nitzschia spp.), dinophytes and the chroococcalean cyanobacterium Synechococcus sp. The nutrient enrichments had no effect on the total biomass of N₂-fixing cyanobacteria. Nevertheless, the biomass of Anabaena spp. was highest in the enrichments with a low N/P ratio. Chlorophyll a concentration and total phytoplankton biomass were not affected by Fe or EDTA, but Fe alone had a positive effect on the chlorophyte Kirchneriella sp. The N₂-fixing cyanobacteria Aphanizomenon sp. responded positively to Fe alone and to both Fe and EDTA added together. The hepatotoxin concentration increased during the experiment, but no clear responses to nutrient enrichments were found. Our study showed species-specific responses to nutrient enrichments among the N₂-fixing cyanobacteria. Although the total phytoplankton production was not Fe-limited; the availability of Fe clearly affected the phytoplankton community structure.     

哈巴湖国家级自然保护区油蒿群落生态化学计量特征对群落生物量和物种多样性的影响

以宁夏哈巴湖国家级自然保护区内的油蒿群落为研究对象,测定了油蒿群落不同发育阶段植物叶C、N、P含量,分析了植物叶C∶N∶P比化学计量特征,探讨了油蒿群落生态化学计量特征对物种多样性和生物量的影响。结果表明,在哈巴湖国家级自然保护区,油蒿群落发育早期,植物生长受N限制,发育中后期,植物生长受P限制;油蒿群落发育早期阶段的植物具有较高的叶C∶N比和较低的叶N∶P比,而油蒿群落发育后期阶段的植物具有较低的叶C∶N比和较高的叶N∶P比。随着叶C∶N比的增大,油蒿群落生物量呈指数函数显著降低,物种多样性呈对数函数降低的趋势,但不显著;而随着叶N∶P比的增大,油蒿群落生物量和物种多样性均呈幂函数显著增加,表明生态化学计量特征对油蒿群落的物种多样性和生物量有重要影响。     

PHOSPHORUS,NITROGEN AND THE GROWTH OF ALGAE IN LAKE KINNERET1

The intracellular concentrations of carbon, nitrogen, phosphorus and chlorophyll a of phytoplankton and zooplankton in Lake Kinneret were determined from 1969 to 1973. The ratios C:P, C:N, chlorophyll a:P, chlorophyll a:N of the algae showed fluctuations which could be related to the nutrient conditions that influence the annual pattern of phytoplankton development, especially in respect to the dinoflagellate bloom of Peridinium cinctum (OFM) Ehr. f. westii (Lemm.) Lef. Relatively high intracellurar P values at the start of the bloom indicated adequate availability of this nutrient and luxury consumption over a short period of time. Later, Peridinium continued to grow despite unusually high C:P ratios (> 300:1). In most years, phosphorus may have eventually limited growth, however, in 1970, the bloom censed despite comparatively high intracellular P concentrations. These observations, together with supplementary evidence from nutrient addition experiments and determinations of specific alkaline phosphatase levels, indicated that, for most of the growth phase of the bloom, Peridinium cells were not directly limited by P. The decline of the bloom usually, but not always (e.g., in 1970), was marked by very high C:P ratios. Thus, a shortage of P may often be a contributory factor to the cessation of the Peridinium bloom and may be limiting phytoplankton growth in the fall. Over the years 1969–73, possibly due to an overall drop in salinity, there appears to be a trend to lower levels of biologically bound phosphorus in Lake Kinneret, without a concomitant decrease in carbon biomass.