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.     

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.     

Automated Pattern Recognition of Phytoplankton – Procedure and Results

Pictures of phytoplankton samples were analyzed as raster images by means of a television camera and a Robotron 4200 computer. A feature vector described the objects irrespective of their angle. Each of the five genera involved were identifiable by a characteristic point cluster in a p-dimensional feature space. A learning method was used during development of the classification structure, and the quality of identification was increased incrementally to the greatest possible degree. Asterionella formosa was identified in all cases without error despite the relatively coarse scanning grid. Errors in the identification of Fragilaria crotonensis can be reduced by improving the resolution (over 100 picture elements per colony).     

COMPARATIVE KINETIC STUDIES OF NITRATE-LIMITED GROWTH AND NITRATE UPTAKE IN PHYTOPLANKTON IN CONTINUOUS CULTURE1

A comparative study of nitrate-limited growth and nitrate uptake was carried out in chemostat cultures of Ankistrodesmus falcatus (Corda) Ralfs., Asterionella formosa Hass., and Fragilaria crotonensis Kit. In each species growth rate (μ) was related to total cell nitrogen or cell quota (q) by the empirical Droop growth function. Nitrate uptake was a function of both external N concentration and q. The apparent maximum uptake rate (V_m') at a given μ was inversely related to q – q₀, where q₀ is the minimum quota. The apparent half-saturation constant for uptake, (K_m') appears to show a slight inverse trend with μ, although statistical analysis shows that this trend is inconclusive. When q approaches q₀, V_m' is several orders of magnitude greater than μq, the calculated steady-state uptake rate. As q increases, however, the difference between these two variables decreases sharply until q approaches q_m, the cell quota for nitrogen-rich cells. At this point the difference between μq and V_m' disappears. This behavior is explained by the feedback regulation of N uptake. The inverse relationship between V_m' and q – q₀ can be described by an empirical three-parameter equation.     

Untersuchungen zum Phosphatmetabolismus einer Trinkwassertalsperre

The examinations showed that the phosphate supply for the planktonic primary production comes almost exclusively from the influxes. The nutrient pool in the hypolimnion and in the sediment does not play a leading part. Evidently, that is often so in reservoirs. Both the phytoplankton (the most frequent species are Asterionella formosa and Fragilaria crotonensis) and the concentration of phosphate fractions are subject to considerable seasonal variations. The Asterionella spring maximum, typical of this reservoir, is released directly or indirectly by meteorological factors. An adequate orthophosphate concentration is a basic condition which is accomplished, however, nearly every year. The autumnal Fragilaria maximum begins when a definite mixing depth hae been achieved. The breakdown of a diatom maximum is caused almost at all times by exhaustion of the phosphate reserves.     

Natural community bioassays to determine the abiotic factors that control phytoplankton growth and succession

SUMMARY. 1. The successional patterns of the dominant phytoplankton species in Lake Maarsseveen (The Netherlands) were very similar in 1980 and 1981. In December/January the diatoms Stephanodiscus hantzschii Grun., Stephanodiscus astraea (Ehr.) Grun. and Asterionella formosa Hass. dominated the algal community (A. formosa had several further population increases during the year). Fragilaria crotonensis Kitt. came to dominance in March/April, followed by the chrysophyte Dinobryon divergens Imhof and the diatom Cyclotella comta (Ehr.) Kütz in May/June. A second appearance of D. divergens was observed in July/August, followed in 1980 by F. crotonensis and a third small increase of D. divergens. In both years S. astraea and S. hantzschii started to grow again in November/December. Cryptophyceae and Chlorophyta were present throughout the year, but did not show a distinct succession. 2. Natural community bioassays, performed under natural light and temperature conditions in a newly developed outdoor bioassay apparatus, showed that phosphate was the major nutrient limiting the growth rate of the phytoplankton. From January till June, during the decline in phosphorus concentration, the diatoms became successively phosphate limited in the sequence: S. hantzschii, S. astraea, F. crotonensis, A. formosa and C. comta. Light limitation was probably the major cause of the relatively late start of F. crotonensis in early spring. 3. D. divergens, increasing after the diatoms from June till September, was stimulated by the addition of a chelator (EDTA). The chelator might stimulate the formation of trace metal species favouring their uptake (e.g. iron). 4. The patterns of succession of the diatoms observed from January till June and from July till December were to a large extent symmetrical. The controlling factors followed opposite trends: declining phosphorus concentrations with increasing irradiance from winter till spring and increasing phosphorus concentrations with decreasing irradiance from summer till late winter.     

Seasonal light characteristics for a newly formed reservoir in Southwestern USA

Seasonal transparency and extinction coefficients are presented for two major habitats in the Lake Powell system, a large newly formed reservoir in the arid southwestern USA. These habitats include a shallow side canyon station subjected to periodic influx of dissolved and suspended materials and an open bay system. Maximum Secchi disk levels in the open bay station were at 7.7 meters with the Secchi disk estimated to disappear at a level of penetration approximating 10% of the solar radiation entering the lake surface. Mean compensation depth was calculated at 11.7 meters. Seasonal extinction coefficient for unfiltered light in the deeper bay station was 0.321 while extinction coefficients for blue, green and red filtered light were 0.323, 0.309 and 0.474 respectively. The effect of periodic influx of dissolved and suspended substances on the underwater light climate and phytoplankton populations are discussed for desert lakes.This study was supported in part by funds provided by the Navajo and Kaiparowits participants.This study was supported in part by funds provided by the Navajo and Kaiparowits participants.     

Limnological effects of peatland drainage and fertilization as reflected in the varved sediment of a deep lake

Stratigraphy of diatoms and chemistry in the surface sediment deposited at 35 m depth in Lake Polvijärvi was studied. The existence of annual laminations or varves in the sediment allowed a precise dating of the profile. Diatoms were analysed in 0.5 cm sequences; from 0 to 16.0 cm continuously and then intermittently every fourth 0.5 cm down to 44.0 cm. Sediment chemistry (loss-on-ignition, C, N, Fe, Mn, Mg, P, chlorophyll and carotenoids) was analysed from sediment surface down to 10.5 cm of altogether 33 subsamples, each containing 1–3 varves, and spanning the period 1921–1980. From 4.5 cm depth upwards the diatom concentration strongly increases, and the plankton diatom succession from Tabellaria flocculosa through Asterionella formosa to Melosira ambigua and Fragilaria crotonensis reflects a marked eutrophication of the lake. This algal succession occurs in pace with an increase in sediment accumulation rate and changes in sediment chemistry, which indicate increased allochthonous inputs and enhanced algal production in the lake. The change of the lake ecosystem is contemporaneous with extensive peatland draining and fertilizing that was carried out on its watershed during the past two decades. Existing chemical data from a number of lakes situated within the drainage area prove that at present the treated peatlands are the main source of nutrient loading of Lake Polvijärvi. A former period with indications of slightly increased productivity of the lake was dated by varve counting to AD 1690–1910 (35–12 cm). This period (characterised by Asterionella formosa) may coincide with that of the slash-and-burn cultivation in the area.     

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.     

Transplant experiments. Growth-limiting factors for diatoms and bluegreen algae in Norwegian lakes

Population densities and total phosphorus concentrations in samples from different lakes of south-eastern Norway were determined. In addition some transplant experiments with dilute phytoplankton populations were carried out. A laboratory batch culture method was used.The diatoms studied may be divided into three ecological groups based on their cell densities and total phosphorus concentrations in the samples. This classification was supported by the experimental results. Cyclotella spp., Asterionella formosa and Tabellaria fenestrata did not grow or had low growth rates above pH 9. Synedra cf. acus and Fragilaria crontonensis had often high growth rates within the pH 9–10 range, but were not able to grow at pH values above 10. High pH-values had no effect on the growth rate of Oscillatoria. Oscillatoria, Synedra and Stephanodiscus were severely growth-limited in filtered water from oligotrophic lakes. Maximum growth rates of all the populations studied were often obtained after addition of phosphate and chelated iron (FeEDTA) in combination to filtered water samples from oligotrophic/mesotrophic lakes.     

Studies on Gull Lake, Michigan

The seasonal and depth distribution of phytoplankton in Gull Lake, a moderately large, temperate, dimictic, recreational lake in southern Michigan, U.S.A., is described and discussed. Diatoms (Fragilaria crotonensis, Cyclotella michiganiana, Asterionella formosa, and Synechococcus) grow mainly in summer. Diatoms grew under the winter ice, and objections are lodged to the classic explanation of seasonal growth in temperate lakes, and to a recent hypothesis concerning the `paradox of the plankton'. Two raphe-bearing large diatom species were considered to be truly planktonic. A Synechococcus species was similar to that described previously only from Loch Leven.     

The influence of carbon dioxide-depletion on growth and sinking rate of two planktonic diatoms in culture

Growth in relation to CO₂-depletion and CO₂-enrichment was investigated for the freshwater diatoms Asterionella formosa and Fragilaria crotonensis in batch cultures. Algal concentration and pH were measured during growth cycles, and inorganic carbon quantities determined by potentiometric Gran titrations and from pH-alkalinity relationships. After the primary growth with CO₂-depletion and pH increase, successive CO₂-enrichments induced further such cycles and produced a final three- to fivefold increase in algal biomass over that of unenriched controls. The extent of CO₂-depletion, and pH rise, was greater in later cycles, indicative of some cellular adaptation. Values of pH reached 9·7 for Asterionella and 9·9 for Fragilaria. The lowest residual quantities of free CO₂ were 0·1 and 0·03 μmol 1^-1 for Asterionella and Fragilaria respectively, which were less than 0·05% of the corresponding residual quantities of total CO₂. The primary limitation of CO₂-uptake and growth was probably related to the concentration of free CO₂, given the relative excess of other major nutrients (N, P, Si) in he media used. Limited of CO₂-uptake could be restored without CO₂ additions if the CO₂ present was redistributed between its several forms (increasing free CO₂) by the addition of strong acid, although growth was still restricted.

Limitation of CO₂-uptake, either by CO₂-depletion or the addition of an inhibitor of photo-synthesis (DCMU), increased the sinking rate of Asterionella cells from 0·3 to 1 m day^-1. The possible ecological implications of CO₂-pH-growth and CO₂-pH-buoyancy relationships are discussed, which may contribute to the frequent paucity of diatoms during summer in manv productive lakes.     

The Effect of Fish Introductions on the Diatom and Cladoceran Communities of Lake Opeongo,Ontario, Canada

Fish introductions are one of the most widespread anthropogenic perturbations to aquatic ecosystems. Paradoxically, the effects of these introductions on aquatic ecosystems are typically poorly documented. This project studied the effect of fish introductions on Lake Opeongo, an oligotrophic lake in Algonquin Provincial Park, Ontario, Canada (45° 42′ N, 78° 22′ W), using the remains of algae (diatoms) and zooplankton (cladocerans) preserved in the sediments. It was hypothesized that the introduction of cisco or lake herring (Coregonus artedii Lesueur) in 1948, which filled the underutilized pelagic forage fish niche, should have altered nutrient availability for phytoplankton. Prior to cisco introduction, the diatom community of Lake Opeongo reflected a relatively stable oligotrophic state established before European settlement, and consisted of the Cyclotella stelligera complex with subdominants Tabellaria flocculosa IIIp and the Aulacoseira distans complex. No marked changes occurred until ca. 1962 when the diatom community shifted to an assemblage with increased total phosphorus preferences, consisting of Asterionella formosa and lesser amounts of Cyclotella bodanica var lemanica, the C. stelligera complex, Fragilaria crotonensis and T. flocculosa IIIp. The dominant cladoceran Bosmina longirostris increased significantly in relative abundance since the introduction of cisco. The most likely cause of this shift was increased nutrient recycling and/or trophic level changes caused by human manipulation of the fish community of the lake.     

PHOSPHATE AND SILICATE GROWTH AND UPTAKE KINETICS OF THE DIATOMS ASTERIONELLA FORMOSA AND CYCLOTELLA MENEGHINIANA IN BATCH AND SEMICONTINUOUS CULTURE1

Information on the nutrient kinetics of Asterionella formosa Hass. and Cyclotella meneghiniana Kutz. under either phosphate or silicate limitation was obtained for use in a Monod model and in a variable internal stores model of growth. Short-term batch culture growth experiments were fit to the Monod model and long-term semicontinuous culture experiments and short-term uptake experiments were fit to the variable internal stores model. Mathematical analysis indicates that the parameters of the 2 models may be expressed in terms of each other at steady state. The qualitative results of both batch and steady state culture methods agree. For limiting phosphate experiments. A. formosa is better able to grow at low PO₄-P concentrations than C. meneghiniana, as shown by its lower K for PO₄-P limited growth. The k_Q of A. formosa compared to C. meneghiniana found in long-term semicontinuous culture indicates that A. formosa is almost an order of magnitude more efficient at using internal phosphate for growth. The qualitative results under silicate-limited growth of C. meneghiniana is less than that of A. formosa. The k_Q from semicontinuous culture experiments indicates that C. meneghiniana is the more efficient at using internal silicate for growth. Nutrient uptake experiments showed more variability from a Michaelis-Menten relationship than short-term growth experiments. There were no significant differences between the 2 species in half saturation constants for either phosphate or silicate uptake. We observed a marked dependence of the coefficient of luxury consumption (R) of phosphate on the steady state growth rate. A. formosa has a higher R than C. meneghiniana.     

Erratum to: How do UV radiation, temperature, and zooplankton influence the dynamics of alpine phytoplankton communities?

Plankton in mountain lakes are confronted with generally higher levels of incident ultraviolet radiation (UVR), lower temperatures, and shorter growing seasons than their lower elevation counterparts. The direct inhibitory effects of high UVR and low temperatures on montane phytoplankton are widely recognized. Yet little is known about the indirect effects of these two abiotic factors on phytoplankton, and more specifically whether they alter zooplankton grazing rates which may in turn influence phytoplankton. Here, we report the results of field microcosm experiments that examine the impact of temperature and UVR on phytoplankton growth rates and zooplankton grazing rates (by adult female calanoid copepods). We also examine consequent changes in the absolute and relative abundance of the four dominant phytoplankton species present in the source lake (Asterionella formosa, Dinobryon sp., Discostella stelligera, and Fragilaria crotonensis). All four species exhibited higher growth rates at higher temperatures and three of the four species (all except Dinobryon) exhibited lower growth rates in the presence of UVR versus when shielded from UVR. The in situ grazing rates of zooplankton had significant effects on all species except Asterionella. Lower temperatures significantly reduced grazing rates on Fragilaria and Discostella, but not Dinobryon. While UVR had no effect on zooplankton grazing on any of the four species, there was a significant interaction effect of temperature and UVR on zooplankton grazing on Dinobryon. Discostella and Dinobryon increased in abundance relative to the other species in the presence of UVR. Colder temperatures, the presence of zooplankton, and UVR all had consistently negative effects on rates of increase in overall phytoplankton biomass. These results demonstrate the importance of indirect as well as direct effects of climate forcing by UVR and temperature on phytoplankton community composition in mountain lakes, and suggest that warmer climates and higher UVR levels may favor certain species over others.     

Eutrophication and recovery of Lake Vesijärvi (south Finland): Diatom frustules in varved sediments over a 30-year period

Lake Vesijärvi was loaded by sewage from the City of Lahti for 60 years until 1976 when the discharge was diverted. Paleolimnological analyses of the varved bottom sediment indicate that the sedimentation rate within the Enonselka basin, the most eutrophic part of the lake, has been as high as 2 cm yr^–1, and total phosphorus accumulation was 20–40 g P m^–2 yr^–1, during the last 20 years. Within the less eutrophic Laitialanselkä basin, the sedimentation rate did not exceed 1 cm yr^–1, and the formation of varved sediment only began at the end of the 1960's, i.e. about 10 years later than in Enonselkä.Planktonic diatom production was highest in the Enonselka basin. The most abundant diatoms in the sediment between 1970–1985 were Asterionella formosa, Aulacoseira islandica and Stephanodiscus spp. Fragilaria crotonensis and Tabellaria fenestrata had low abundances in the middle of the 1970's but increased again at the end of the 1970's. Asterionella formosa and Diatoma elongatum reached their maxima between 1979–1984 when the hypolimnion of the Enonselk/:a basin was aerated artificially. In the Laitialanselkä basin, the production of planktonic diatoms has been lower and the species composition of the diatom community differed from that in Enonselkä. However, at the end of 1980's the total accumulation of diatoms in Laitialanselkä approached levels which were observed at the end of 1950's in Enonselkä, prior to the rapid eutrophication period.The production and thereby the sedimentation of diatoms has decreased towards the end of the 1980's in Enonselkä, indicating reduced nutrient availability in the lake water. This reduction was due to the decreased external loading of phosphorus as well as to the decreased release of phosphorus from the sediment as a result of improved oxygen balance in the hypolimnion.     

CHANGES IN THE PLANKTONIC DIATOM FLORA OF A LARGE MOUNTAIN LAKE IN RESPONSE TO FERTILIZATION1

Fertilization (phosphorus and nitrogen) of Kootenay Lake, British Columbia, was begun in 1992 in an attempt to increase food resources for declining stocks of kokanee salmon. In 1992 and 1993, significant changes in diatom abundance, species composition, diversity, and morphology of individual species were observed in the fertilized North Arm in comparison to the unfertilized South Arm. Effects of fertilization were most obvious during the spring. During summer stratification, fertilization did not result in significant changes in total abundance, but changes in species composition were observed. In 1993, with continued fertilization of the North Arm, diatom abundance in both spring and summer was significantly higher than at the same sites in 1992. No significant differences were observed in the unfertilized South Arm. Diatom abundance was significantly correlated with dissolved inorganic nitrogen (positive) and dissolved reactive silica (negative). Diatom assemblages in the fertilized North Arm in both the spring and summer of 1993 were dominated by Asterionella formosa and Fragilaria crotonensis, taxa considered mesotrophic. Abundance of the oligotrophic taxa Rhizosolenia eriensis and Cyclotella michiganiana was higher in the South Arm than in the North Arm. Differences in diatom assemblages between the two arms in both years were further summarized by multivariate statistical analyses. Morphological variations in two small Cyclotella species (C. comensis and C. cf. pseudostelligera) were observed between the fertilized and “control” stations. Populations of both species from fertilized stations exhibited stronger silicification with coarser striations and a smaller mean size.     

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.     

Spatial heterogeneity of diatom silicification and growth in a eutrophic reservoir

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