TEMPERATURE EFFECTS ON SILICON- AND PHOSPHORUS-LIMITED GROWTH AND COMPETITIVE INTERACTIONS AMONG THREE DIATOMS1

Three diatom species, Stephanodiscus hantzschii (Ehr.) Grun., Asterionella formosa Hass. and Fragilaria crotonensis Kitt. Hass. were isolated from Lake Maarsseveen where they are dominant and show a successional sequence. The physiological responses of each species to temperature and limitation by silicon and phosphorus were determined over the temperature range of 5° to 20° C using short-term batch culture methods. Stephanodiscus hantzschii had higher maximum growth rates than the other two species at all temperatures, and the maximum growth rates of all species increased with increasing temperature. Temperature affected not only maximum growth rates but also half-saturation constants (K_s) and the minimum cell quotas. S. hantzschii had low silicon requirements for growth under Si-limiting conditions, and A. formosa and F. crotonensis had higher and nearly identical silicon requirements. The K_s values for silicon for S. hantzschii were essentially constant from 5° to 20° C but varied greatly for the other two species. A. formosa had the lowest requirements for growth under phosphorus limitation, F. crotonensis was intermediate and S. hantzschii had the highest growth requirements for phosphorus. The K₁ values for phosphorus were constant over the temperature range for both A. formosa and F. crotonensis and were much higher and variable for S. hantzschii. Nutrient competition experiments were performed in continuous cultures at four temperatures and various Si:P ratios. The results generally, but not always, confirmed the predictions based on the Monod relationships for each species. Results not in agreement with predictions were usually because of similar physiological properties of A. formosa and F. crotonensis or because of decreased loss rates for F. crotonensis due to wall growth. In cultures with all three species phosphorus-limited (Si:P > 75), A. formosa often dominated as predicted, although F. crotonensis was sometimes the most abundant species. As predicted, S. hantzschii never dominated at high Si:P ratios. At intermediate Si:P ratios when A. formosa and F. crotonensis were both Si-limited and S. hantzschii P-limited, all three species coexisted because A. formosa and F. crotonensis have almost identical silicon requirements, although sometimes F. crotonensis was more abundant than predicted. At 10°C the results agreed best with the predictions; A. formosa dominated at high Si:P ratios and S. hantzschii dominated as predicted at low Si:P ratios when all three species were Si-limited.     

The recent diatom history of Loch Leven, Kinross

Diatoms in the top metre of lake sediment have been studied for indications or recent changes in the environment of Loch Leven. The diatoms indicate a eutrophic, alkaline lake with Melosira italica ssp. subarctica, Tabellaria Flocculosa and Stephanodiscus astraea the dominant taxa at the base of the 1-m section. There is then an appeance of Melosira ambigua and Stephanodiscus hantzschii suggesting a richer environment. Post A.D. 1816 changes in taxa suggest possible correlation with the lowering of lake level in 1830 and the dominance of Diatoma elongatum, Cyclotella pseudostelligera, Stephanodiscus astraea var. minutula, S. tenuis, S. hantzschii, and Synedra rumpens and var. soctica in zone 4 agrees with present-day plankton records.     

Das Sediment des Kummerower Sees. Untersuchungen des Chemismus und der Diatomeenflora

The Sediment of Lake Kummerow. Investigations on the Chemism and the Diatom Flora The paper deals with the distribution of chemical parameters and of diatoms in the deposits of the eutrophic Lake Kummerow (GDR), produced during about the last 3500 years. The sediment consists of a calcareous gyttja with a very low content of sand and clay. The results indicate a relatively constant trophic state of the lake in former times and an increased production within the last five centuries. Stephanodiscus astraea and its variety minutula is by far the most constant diatom species; in the uppermost parts of the sediment small sized planctonic taxa are increasing, in the main Stephanodiscus hantzschii var. pusillus, Melosira granulata var. angustissima, Asterionella formosa and Synedra acus var. angustissima and var. radians.     

Can Hydrodynamics Change Phosphorus Strategies of Diatoms?—Nutrient Levels and Diatom Blooms in Lotic and Lentic Ecosystems

Diatom blooms occur in many water bodies worldwide, causing significant ecological and social concerns. In order to understand the mechanisms of diatom blooms formation, the effects of varying phosphorus (P) concentration and hydrodynamics on the growth of diatoms were studied by combining results from field observations and laboratory experiments. The field investigation showed that spring diatom blooms (Cyclotella meneghiniana and Stephanodiscus hantzschii) occurred in Lake Taihu and Hanjiang River with similar environmental factors such as water temperature, pH, and dissolved oxygen in 2008. Concentrations of total phosphorus (TP), total nitrogen, and ammonia nitrogen (NH₄-N) in Lake Taihu were significantly higher than the concentrations in the Hanjiang River. Laboratory experiments were conducted to evaluate growth and physiological responses of four lotic diatoms (Cyclotella atomus, Fragilaria crotonensis, Nitzschia palea, and S. hantzschii, isolated from the Hanjiang River) and three lentic diatoms (C. meneghiniana, Melosira varians, and Stephanodiscus minutulus, isolated from Lake Taihu, Lake Donghu, and Guanqiao Pond, respectively) to various P concentrations under small-scale turbulent and standing conditions. Our results showed that, with turbulence, lotic diatoms C. atomus, F. crotonensis, N. palea, and S. hantzschii demonstrated a significant increase in affinity for P compared with lentic diatoms C. meneghiniana, M. varians, and S. minutulus. Moreover, lotic diatoms C. atomus, F. crotonensis, and N. palea had higher growth rates and photosynthetic efficiencies with small-scale turbulence than with standing conditions both in P-limited and P-replete conditions. Lotic species S. hantzschii and three lentic diatoms (C. meneghiniana, M. varians, and S. minutulus) grew well under standing conditions. Our results may explain our field observation that the occurrence of diatom blooms in lakes is often associated with higher TP concentrations whereas in rivers, diatom blooms occur at a wide range of TP concentrations under flows. Therefore, different hydrodynamics and nutrient concentrations determined the dominant diatom species, according to their habitat-dependent physiological characteristics.     

The influence of irradiance,photoperiod and temperature on the growth kinetics of three planktonic diatoms

The influence of irradiance, photoperiod and temperature was determined for the growth kinetics of the diatoms Aulacoseira subarctica, Stephanodiscus astraea and Stephanodiscus hantzschii and the results compared with those of cyanobacteria. Irradiance and photoperiod relationships were qualitatively similar to those for cyanobacteria in that: (1) growth rate (K) was proportionally greater under short photoperiods, with ratios of K under continuous light to K under 3:21 light:dark (LD) cycles of 1·50, 1·80 and 2·96 for A. subarctica, S. astraea and S. hantzschii respectively; (2) at subsaturating irradiances, K was proportional to irradiance and independent of temperature with a negligible predicted maintenance growth rate requirement. Apparent growth efficiencies (GE) at subsaturating irradiances were 0·26±0·03, 0·42±0·03 and 0·50±0·03 divisions mol^-1m² for A. subarctica, S. astraea and S. hantzschii with the values for Stephanodiscus species comparable to values for Oscillatoria species. Under a 3:21 LD cycle at 4 °C, light-saturated growth rates were 0·066±0·004, 0·197±0·033 and 0·285±0·018 divisions day^-1 for A. subarctica, S. astraea and S. hantzschii. S. hantzschii growth rate at 4 °C exceeded maximum Oscillatoria growth rates at 23 °C and the S. astraea growth rate at 4 °C was equivalent to O. agardhii growth rate at 20 °C. Temperature increases above 4 °C gave Q₁₀ values between 4 °C and 12 °C of 3·68, 2·39 and 1·92 for A. subarctica, S. astraea and S. hantzschii, but higher temperatures resulted in minor increases in K. S. astraea growth rate peaked at 16 °C, declining sharply at higher temperatures. February to March in situ growth rates in Lough Neagh, mean temperature 4·3 °C, showed that the A. subarctica in situ K of 0·058 divisions day^-1 was close to the laboratory K at 4 °C, but that S. astraea in situ K of 0·101 divisions day^-1 was lower than the laboratory K at 4 °C.     

BENTHIC MARINE DIATOM ASSOCIATIONS: UPPER FLORIDA BAY (FLORIDA) AND ASSOCIATED SOUNDS1

Models of the diatom associations found in upper Florida Bay and adjoining sounds have been constructed utilizing Q modal factor-vector analysis and ecologic diversity indices (Shannon index, number of species, evenness). Four distinct associations were defined using Q-mode factor-vector analysis. Two associations were epiphytic, occurring on Thalassia testudinum König—Association I was characterized by Cocconeis placentula Ehr., Association III by Cylindrotheca closterium (Ehr.) Reim. & Lewin and Cocconeis placentula Ehr. The other two associations were epipelic, occurring on the carbonate mud substratum—Association II was characterized by Cyclotella striata (Kütz.) Grun., Rhopalodia gibberula (Ehr.) O. Müller and Surirella fastuosa (Ehr.) Kütz., Association IV by Fragilaria crotonensis Kitton and Cyclotella striata (Kütz.) Grun. The majority of the 161 species identified were present in both the epiphytic and epipelic assemblages. Only 33 species were restricted to the epiphyton, and 18 species restricted to the epipelon. The epipelic assemblage was significantly more diverse than was the epiphytic assemblage. A general trend of increased diversity away from terrestrial environs toward more open areas of water in both the epipelon and epiphyton was also found.     

Succession of phytoplankton in a deep stratifying lake: Mondsee,Austria

Phytoplankton numbers, biovolume, chlorophyll-a and various physico-chemical characteristics were followed at weekly intervals in Mondsee, Austria during the year 1982. Secchi-disk transparency varied from 10 m in winter to 2 m in September. Prior to the onset of stratification phosphate-phosphorus concentration was 4 µg 1^–1 decreasing to undetectable values thereafter. Nitrate-nitrogen dropped from 590 µg 1^–1 to about 100 µg 1^–1 during the same time. The vernal bloom was dominated by Asterionella formosa Hass. which abruptly declined after silicon depletion. Spring growth ceased in early June, when Tabellaria flocculosa (Lyngb.) Kütz var. asterionelloides Grun. dominated. Oscillatoria rubescens D.C. and Microcystis aeruginosa Kütz. dominated summer and early autumn followed by the chrysophyte Dinobryon divergens Imh. and D. sociale Ehr. which formed up to 69% of total biovolume in October. Thereafter diatoms and Cryptophyceae (Rhodomonas lacustris Pascher and Ruttner, Cryptomonas pusilla Bach.) became abundant again.Maximum chlorophyll-a concentration in the epilimnion (16 µg 1^–1) was reached during spring growth of the diatoms. During summer higher chlorophyll-a levels were always associated with the metalimnetic layer of Oscillatoria.Compared with earlier studies, both the total biovolume and the share of Oscillatoria rubescens significantly decreased because of reduced nutrient loading of the lake and wash-out of Oscillatoria (theor. renewal time of the lake: 1.7 years).     

Resource requirements of four freshwater diatom taxa determined by in situ growth bioassays using natural populations from alpine lakes

We performed a series of in situ batch culture experiments to assess the resource requirements of common diatom taxa in alpine lakes of the central Rocky Mountains of North America. While physiological data are available on the resource requirements of some of these taxa, it is unclear whether intraspecific generalizations can be made across aquatic systems due to the potential development of ecotypes. In these experiments, we used amended lake water for a culture medium and natural diatom populations. Growth kinetics were determined for Asterionella formosa Hassall, Fragilaria crotonensis Kitton, Staurosirella pinnata (Ehr.) Williams and Round and Tetracyclus glans (Ehr.) Mills. Staurosirella pinnata, a historically abundant alpine diatom, had very low N and P requirements. Asterionella formosa and F. crotonensis, generally considered meso- or eutrophic species, exhibited low P requirements if N and Si were in moderate supply. Tetracyclus glans had the highest Si requirement. These experiments reveal that the recent changes in diatom community structure in these alpine lakes may be driven by changes in nutrient supply. We suggest that local diatom taxa and a natural culturing medium should be used to obtain more representative algal physiological data from a particular area.     

Ingestion and incorporation of freshwater diatoms by Daphnia pulicaria: do morphology and oxylipin production matter?

In order to explain differences in the growth and reproductionof Daphnia pulicaria fed various freshwater diatoms, we measuredingestion rates and carbon incorporation for six cultured diatomspecies: the single-celled Stephanodiscus hantzschii, Stephanodiscusminutulus and Cyclotella meneghiniana, and the colony-formingAsterionella formosa, Fragilaria capucina and Fragilaria sp.Two of the colony-forming species, when damaged, produced polyunsaturatedaldehydes (oxylipins) that have been found to impair the reproductionof marine copepods. We tested two hypotheses: (i) feeding andincorporation rates are affected by diatom morphology; and (ii)polyunsaturated aldehydes act as feeding deterrents. Daphniabody length versus ingestion rate regressions differed for single-celledand colony-forming diatoms. Ingestion rates for single-celleddiatoms showed clear size dependencies and high correlationcoefficients, while the dependency was weak for colony-formingdiatoms and individual variability was high. This differencewas not observed for carbon incorporation rates, which showedlow variability for all diatoms. Asterionella formosa yieldedthe lowest incorporation rates due to low incorporation efficiency,while all other diatoms were incorporated at similar rates.Thus, morphological differences of the diatoms had no effecton carbon uptake by Daphnia. The presence or absence of polyunsaturatedaldehydes did not cause different ingestion rates; hence thealdehydes are not feeding deterrents.     

The production ecology of Stephanodiscus Astraea (Ehrenb.) Grun

The production ecology of Stephanodiscus astraea (Ehrenb.) Grun. is discussed with respect to other phytoplankton growths, mixing regimes and nutrient availability. Populations of Stephanodiscus were studied during 1972 and 1973 in an artificially mixed reservoir with a capacity of 3.5 × 10⁷m³. Major nutrients such as soluble reactive phosphates (60–200 μg/1) and nitrates (1–8 mg/1) were not limiting. Large amounts of silicate (approximately 10 mg/1) were utilized to support moderately large populations of S. astraea (1.3 × 10⁷ μm³/ml). Light penetration and mixing regimes strongly affected the periodicity and size of standing crops. It is suggested that heterotrophic production can interact with the mixing environment to influence the production of turbid waters.     

Winter blooms of centric diatoms in the River Danube and in its side-arms near Budapest (Hungary)

Although the phytoplankton in large eutrophic rivers is often abundant, data on winter populations are scarce. Phytoplankton has been sampled since 1979 in the main arm of the River Danube at Göd (20 km north from Budapest), and also from two side-arms, upstream and downstream of Budapest. Analysis of winter samples has shown high population densities of phytoplankton blooms on several occasions. In general centric diatoms (20 taxa determined) were abundant and dominant during the blooms. The most abundant taxon in November was Stephanodiscus hantzschii f. tenuis, and in February S. minutulus. The most important factors in forming winter blooms of Centrales are rich nutrient supply, slow current speed and high transparency.Massive phytoplankton blooms occurred several times between November and March in the small side-arm at Göd. In the large side-arm downstream of Budapest (Soroksári-Dung) large blooms of centric diatoms were found in winter (often under ice, which was covered by snow in many cases). Here S. hantzschii f. tenuis and S. minutulus were the most abundant species.     

The effect of fungal parasitism on the succession of diatoms in Lake Maarsseveen I (The Netherlands)

SUMMARY. During late winter and early spring, diatoms are predominant in Lake Maarsseveen I and show a succession in blooming. To elucidate the role of parasitism in this succession, samples were taken once a week in 1978, 1979and 1980 and examined for fungal parasites, further, factors which presumably influenced the phytoplankton growth. including temperature and chemical compounds, were studied. Parasites were found on Asterionella formosa Hass., Stephanodiscus astraea (Ehr) Grun. and Cyclotella comta (Ehr) Kütz. The chytrid fungus Zygorhizidium planktonicum
Canter was parasitic tin Asterionella formosa. This was the only case where high infection percentages were registered. The developmental phases of Zygorhizidium plunktonicum on Asterionella formosa were studied. The fungus was capable of affecting the spring increase and bringing about a premature end to the bloom. Only when the fungus was temporarily inhibited in its parasitic activity (this phenomenon coincided with periods of frost) was A. formosa able to reach a high abundance. Severe parasitism on A. formosa favoured the development of other diatoms (mainly of Fragilaria crotoriensis Kitt., Stephanodiscus hantzschii Grun. and Stephanodiscus astraea) .
Host-parasite interrelationship studies performed in the laboratory showed that temperature can be an important environmental factor associated with epidemics of Zygorhizidium plankiomcum on Asterionella formosa. At very low temepratures (1.5. 1) the fungal activity was inhibited while Asterionella fortnosa still grew well. However, at 5, 10 and 18 the fungus manifested a high infection rate and was able to overtake A sterionella .     

Über Rassenbildung bei der zentrischen KieselalgeStephanodiscus hantzschii Grun.

Zusammenfassung Stephanodiscus hantzschii Grun. ist mit großer Wahrscheinlichkeit keine einheitliche Art, sondern aus mehreren strukturellen Varietäten sowie mehreren Rassen zusammengesetzt, die sich im statistischen Mittel durch ihre Schalendurchmesser sowie durch ökologische Ansprüche voneinander unterscheiden. Die relative und die nach der Häufigkeit in den untersuchten Gewässern gegebene Verteilung der Schalendurchmesser weist mehrere Gipfel auf, die den Größenschwerpunkten der Rassen gleichgesetzt werden. Jedes der untersuchten mecklenburgischen Gewässer hat ein eigenes, charakteristisches Spektrum von Schalendurchmessern, wobei diese Spektren eine Beziehung zum Nährstoffgehalt (Phosphor und Stickstoff) des Gewässers erkennen lassen.Der technischen Assistentin FrauMarkwardt dankt der Verfasser für ihre gewissenhafte und interessierte Mitarbeit.     

Spatial and temporal dynamics of phytoplankton communities along a regulated river system, the Nakdong River, Korea

The longitudinal distribution and seasonal fluctuation of phytoplankton communities was studied along the middle to lower part of a regulated river system (Nakdong River, Korea). Phytoplankton biomass decreased sharply in the middle part of the river (182 km upward the estuary dam), and then increased downstream reaching a maximum at the last sampling station (27 km upward the estuary dam). In contrast, there was little downstream fluctuation in species composition, irrespective of pronounced differences in nutrient concentrations (TN, TP, NO₃, NH₄, PO₄) as well as in algal biomass. In the main river channel, small centric diatoms (Stephanodiscus hantzschii, Cyclotella meneghiniana) and pennate diatoms (Synedra, Fragilaria, Nitzschia) were dominant from winter to early spring (November–April). A mixed community of cryptomonads, centric and pennate diatoms, and coenobial greens (Pediastrum, Scenedesmus) was dominant in late spring (May–June). Blue-green algae (Anabaena, Microcystis, Oscillatoria) were dominant in the summer (July–September). A mid-summer Microcystis bloom occurred at all study sites during the dry season, when discharge was low, though the nutrient concentration varied in each study site. Nutrients appeared everywhere to be in excess of algal requirement and apparently did not influence markedly the downstream and seasonal phytoplankton compositional differences in this river.     

INTRAGENOMIC NRDNA ITS POLYMORPHISM IN THE STEPHANODISCUS NIAGARAE COMPLEX (BACILLARIOPHYCEAE)

Intragenomic nrDNA ITS variability was examined in representative species of the Stephanodiscus niagarae complex. PCR was performed on DNA extracted from monoclonal cultures, and sequence variability determined using standard cloning techniques. Preliminary data from all species reveal intragenomic polymorphism. ITS sequence data is being used to delineate closely related species of algae and polymorphisms are potentially problematic for phylogenetic reconstruction and identity. We are exploring the extent of polymorphism in other freshwater Thalassiorsiraceae, including Stephanodiscus hantzschii fo. tenuis and Cyclostephanos invisitatus.     

Testing addition of <Emphasis Type="Italic">Pseudomonas fluorescens</Emphasis> HYK0210-SK09 to mitigate blooms of the diatom <Emphasis Type="Italic">Stephanodiscus hantzschii</Emphasis> in small- and large-scale mesocosms

An algicidal bacterium Pseudomonas fluorescens HYK0210-SK09 (SK09) was applied to a natural bloom of Stephanodiscus hantzschii using a small-scale mesocosm (SM) and a large-scale mesocosm (LM) to clarify the algicidal effects and evaluate the response of the planktonic community and environments. When SK09 cells were inoculated at a final concentration of 5 × 10⁶ cells mL⁻¹, the abundance of S. hantzschii decreased significantly by 95% in SM and 85% in LM. The microcosm in the laboratory revealed that the abundance of Pseudomonas increased rapidly after inoculation with a corresponding decrease in the S. hantzschii population. Nutrient concentrations increased following the decline in the diatom cells. The abundances of nondominant species such as Chlamydomonas, Cryptomonas, and Navicula increased slightly with increased nutrient concentrations. The abundance of heterotrophic protists also increased significantly due to utilization of SK09 as food. The present study demonstrates that SK09 is an effective biocontrol agent for natural S. hantzschii bloom, and grazing pressure plays a crucial role in the successful application of algicidal bacteria to natural environments.     

Effects of algal food quality on sexual reproduction of Daphnia magna

The objective of our study was to investigate sexual reproduction of Daphnia magna associated with mating behaviors and hatching rates, according to different algal food sources. Since a diatom is known to contain more abundant long‐chain poly unsaturated fatty acids (PUFAs), we hypothesized that the diatom‐consuming D. magna would exhibit more successful reproduction rates. Upon the hypothesis, we designed three experiments using two algal species, a green alga (Chlorella vulgaris) and a diatom (Stephanodiscus hantzschii). From the results, we found that the mating frequency and copulation duration increased in the treatment with S. hantzschii, resulting in a significant increase of hatching rates of resting eggs. In the other two repetitive mating strategies (e.g., one female vs. multiple males, and one male vs. multiple females), we found that the hatching rates of resting eggs were greater in the S. hantzschii treatment. In addition to the mating strategy, male body size significantly increased in the diatom treatment, hence average diameter of penis was also statistically different among the treatments (greater diameter in the S. hantzschii treatment). To examine the effect of algal food quality, we estimated quantity of fatty acids in the two algal species. Our result showed that S. hantzschii had a higher proportion of long‐chain PUFAs than C. vulgaris. Furthermore, a stable isotope analysis revealed that carbon and nitrogen originated from S. hantzschii were more assimilated to D. magna. In summary, our study manifested that diatom consumption of D. magna leads to more successful sexual reproduction. We then discussed how the diatom consumption of zooplankton influences food web dynamics in a freshwater ecosystem.     

Diatom Succession and Interaction in Littoral Periphyton and Plankton

Periphyton and plankton samples were collected at four littoral stations in a relatively shallow, eutrophic lake (Elk Lake, B.C., Canada) over a six month period from August 1967 to January 1968. The most abundant planktonic diatoms demonstrated a pattern of seasonal succession typical of temperate lakes, and all were present in the periphyton. This pattern was identical at all stations:Fragilaria crotonensis was dominant from August to October,Asterionella formosa in November and December, andMelosira italica (plusM. varians) in January.F. virescens, although never dominant, peaked in October. Periphyton communities were dominated byAchnanthes minutissima, Cocconeis placentula, F. crolonensis andF. virescens. Considerable station variation in successional patterns occurred over different exposure periods; station differences were least in those samples immersed for monthly intervals, and greatest in those immersed for a cumulative four-month period. Interaction between the phytoplankton and periphyton was illustrated by the occurrence of species common to both habitats. A decrease in cell numbers and percent abundance of these species in planktonic populations coincided with their increase in the periphyton, a relationship which appeared dependent on the breakdown of thermal stratification in November. For example, following turnover,F. crotonensis andA. formosa settled out of the plankton and correspondingly increased in percent abundance in the periphyton. This interdependence was less evident in the four-month samples, whereA. minutissima andC. placentula dominated throughout the entire period and appeared to out-compete the more typical planktonic components for diminishing substrate area. Species interaction or competition was accentuated as exposure duration and periphyton total cell standing crops increased and species diversity decreased, and appeared to account in part, for station differences in successional patterns.     

Sinking rates of planktonic diatoms in an unstratified lake: a comparison of field and laboratory observations

SUMMARY. 1. Natural population sinking rates were calculated by fitting an exponential regression to 6 years'observations on declining crops of Melosira italica subarctica O. Müll. and Stephanodiscus astraea (Ehr.) Grun.
2. Losses were described by an exponential model which yielded still water sinking rates of 0.86 md⁻¹ for M. italica and 0.45md⁻¹ for S. astraea.
3. Laboratory measurements of the sinking rate of natural populations showed that the rate increased abruptly from less than 0.2 m d⁻¹ in growing populations to 0.4 m d⁻¹ after silica depletion.
4. The measured sinking rate of S. astraea agreed well with that observed in the field. Where as populations of M. italica appeared to sink more quickly in the lake than could be accounted for by laboratory observations.