Factors influencing phytoplankton growth and succession in Farmoor Reservoir

Weekly observations were made at the Thames Water Authority's Farmoor reservoir during the development of the spring bloom of phytoplankters from November 1973 to March 1974. Counts were made of the phytoplankters, their fungal parasites and the zooplankters. Analyses were made of those chemical parameters which it was thought might exert an influence on phytoplankton growth. Physical and meteorological measurements were also made. The development and decline of four phytoplankton populations were observed. The growth of Asterionella formosa Hass. was interrupted in December when 43·5% of the cells were parastiized by Zygorhizidium affluens Canter. Stephanodiscus astraea (Ehr.) Grün. and Stephanodiscus hantzschii Grün. were not similarly affected. All three diatoms were apparently limited finally by silica depletion. The desmid, Closterium peracerosum Gay. became 66% parasitized by an undescribed chytrid and declined rapidly. Light, turbulence, zooplankton grazing and nutrients other than silica appeared to be unimportant.     

The spring development of phytoplankton in Lake Erken

SUMMARY. In Lake Erken there is a regular phytoplankton peak at ice-breakup in the spring. Pechlaner (1970) studied the spring outburst of 1960. We carried out a similar study in 1973, when the climatic factors were completely different, with only 10 cm of ice and almost no snow. Thus, due to light penetration, there was a high phytoplankton biomass under the ice as early as March.
Stephanodiscus astraea (Ehr.) Grun. and Asterionella formosa Hass. dominated the biomass during the spring development in 1973 instead of the normally dominating S. hantzschii v. pusillus Grun.
The nutrient decrease was slower in 1973 than in 1960. A comparison of carbon assimilation and the decrease of nitrate and phosphate indicate sufficient nutrient supply in 1973. In 1960, the population grew logarithmically until the nutrients became limiting. In 1973, the average biomass was greater, but the mean probable primary production was only half that of 1960. From the activity coefficients for the 2 years it is also clear that, in 1960, S. hantzschii v. pusillus was highly productive during its short but intensive growth period. Its minute size and high surface/volume ratio enabled it to take up nutrients rapidly and efficiently. The chlorophyll content of S. astraea in 1973 was only half as much as that of S. hantzschii v. pusillus in 1960. Since the light conditions under the ice in March 1973 were more unfavourable than during the exponential phase of 1960, production was much lower.     

Population dynamics of three planktonic diatoms in lake Constance

A population dynamics analysis for planktonic diatoms is presented that allows estimates of the net rate of increase (k), the death rate (δ), the sedimentation rate (σ) and, in absence of grazing, the growth rate (μ). It requires counts of live and dead cells suspended in the euphotic part of the water column and accumulated in sedimentation traps. The application of the model is demonstrated for the three dominant summe diatom species in Lake Constance. Asterionella formosa Hass, Fragilaria crotonensis Kitton and Stephanodiscus binderanus Krieger. Only during the first two weeks of the summer bloom of diatoms the loss rates were unimportant in comparison to the growth rates. Thereafter diatom population dynamics was strongly influenced by sedimentation and mortality, which sometimes led to a decrease in population density even when cell division continued at high rates. There were two periods of extraordinarily high death rates, which were associated in the case of A. formosa with silicon depletion and in the case of F crotensis with fungal parasitism.     

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.     

Competition between the Cyanobacterium Limnothrix redekei and Some Spring Species of Diatoms under P-Limitation

The nutrient saturated growth and the competition between Limnothrix redekei and some spring diatoms for phosphate were studied in semi-continuous cultures at 10°C with a 6/18 h light/dark cycle and at 15 °C with a 12/12 light/dark cycle. Under these conditions in unialgal cultures, the maximum specific growth rate of Limnothrix redekei was two to three time lower than that of the diatoms. However, Limnothrix redekei was the superior competitor for phosphate under both conditions in mixed cultures. The diatoms in mixed cultures from each species with Limnothrix redekei were competitively displaced but at a different rate. This allowed the diatoms to be ranked in terms of their competitive ability for phosphate at 10°C as follows: Synedra acus > Asterionella formosa > Stephanodiscus minutulus > Nitzschia acicularis > Cyclotella meneghiniana > Stephanodiscus neoastrea > Stephanodiscus hantzschii. The competitive ability of Limnothrix redekei at 15 °C was increased and the rank order of diatoms slightly changed. The hypothesis that cyanobacteria are promoted by phosphate because they are inferior competitors for phosphate is incorrect for Limnothrix redekei.     

The parasitic chytrid, Zygorhizidium, facilitates the growth of the cladoceran zooplankter, Daphnia, in cultures of the inedible alga, Asterionella

In food-web studies, parasites are often ignored owing to their insignificant biomass. We provide evidence that parasites may affect trophic transfer in aquatic food webs. Many phytoplankton species are susceptible to parasitic fungi (chytrids). Chytrid infections of diatoms in lakes may reach epidemic proportions during diatom spring blooms, so that numerous free-swimming fungal zoospores (2-3 microm in diameter) are produced. Analysis shows that these zoospores are rich in polyunsaturated fatty acids and sterols (particularly cholesterol), which indicates that they provide excellent food for zooplankters such as Daphnia. In life-table experiments using the large diatom Asterionella formosa as food, Daphnia growth increased significantly in treatments where a parasite was present. By grazing on the zoospores, Daphnia acquired important supplementary nutrients and were able to grow. When large inedible algae are infected by parasites, nutrients within the algal cells are consumed by these chytrids, some of which, in turn, are grazed by Daphnia. Thus, chytrids transfer energy and nutrients from their hosts to zooplankton. This study suggests that parasitic fungi alter trophic relationships in freshwater ecosystems and may be the important components in shaping the community and the food-web dynamics of lakes.     

Seasonal Distribution of Phytoplankton in Yellowknife Bay,Great Slave Lake

The factors influencing the seasonal distribution of phytoplankton were determined between July 1975 and December 1977 in Yellowknife Bay, Great Slave Lake. The initiation of the spring bloom occurred during April of each year in response to changing light conditions and despite the 0 °C water temperatures. Although the plankton showed some fluctuations in density (100–400 mg/m³) during the summer, there was no vernal decrease in the size of the populations. The end of the growing season was characterized by a sharp drop in numbers near the end of October. Multiple regression analysis indicated that changes in the standing crop of the plankton during the summer were primarily related to temperature (r = 0.60) and the concentrations of NO₃–N (r=–0.74) and SiO₂ (r=–0.82). Changing light conditions probably had little effect on the population during the growing season. Chlamydomonas lapponica was dominant during April of each year. While the growth of this species caused a reduction in the concentrations of total phosphorus and NO₃–N, the level of SiO₂ remained constant and consequently diatoms (Asterionella formosa, Stephanodiscus astraea, Diatoma tenue var. elongatum and Melosira islandica) waxed rapidly during May. By the first of June, nutrient concentrations were low, thereby permitting the development of Dinobryon bavaricum and Dinobryon cylindricum. The extremely rapid decrease in the densities of common species during the fall was probably due to the rapid decline in temperature and light levels and the formation of ice on the bay.     

OPINION Endless summer: internal loading processes dominate nutrient cycling in tropical lakes

SUMMARY. 1. Fossil diatom assemblages deposited in more than a dozen African lakes roughly 9500 years BP were dominated by a single planktonic species, Stephanodiscus astraea (Ehrcnb.) Grun. (although realistically this is likely to be a species complex). These diatoms flourished when lake-levels were maximal. Data are included from many of (he large African lakes, and others extending from Lake Abhé0, Ethiopia, to Lake Cheshi, Zambia.
2. Because the ecological physiology of Stephanodiscus species is well known one can predict the nutrient regime that must have existed when Stephanodiscus bloomed. Owing to competition for resources Stephano-discus species dominate when the supply ratio of silicon to phosphorus (in moles) in the epilimnion is relatively low (Si:P∼1). Consequently, lakes dominated by S. astraea are often hypereutrophic.
3. We propose a series of hypotheses to explain why tropical lakes have decreasing Si:P ratios as lake-levels increase, primarily owing to internal P-loading processes in the epilimnia. These observations appear to contradict present conceptions of the fundamental relationships governing nutrient loadings to and within lakes. Tropical lakes appear to have had increasing epilimnetic phosphorus loading as lake-levels increased. In contrast, large, deep lakes in the temperate zone are usually oligotrophic, with high Si:P ratios.
4. Our major conclusion is that regeneration rates are greater than removal rates for phosphorus in tropical lakes as compared to temperate lakes, especially where epilimnelic mixing exceeds 50 m. Biological control of the elemental cycles dominate in tropical lakes, whereas nutrient cycles in temperate lakes are dominated by physical processes for a large part of the year. This results in major differences in the fundamental mechanisms of nutrient regeneration and their relationships to morphometric features of lakes in the two regions.     

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.     

The A/C (Araphidineae/Centrales) ratio in high and low alkalinity lakes in eastern Minnesota

SUMMARY. Diatoms from surface sediment samples from lakes on both sides of a glacial drift border in eastern Minnesota show that Araphidinate diatom genera like Fragilaria, Asterionella and Synedra are more common in lakes in drift with lower CaCO₃ whereas Stephanodiscus and Melosira species are more abundant in those with high CaCo₃. In a lake sediment core from low carbonate drift Fragilaria crotonensis and Asterionella formosa increased when pioneer farmers arrived. These results confirm speculation that high A/C ratios are associated with low-alkalinity eutrophic lakes.     

HOST PARASITE INTERACTIONS BETWEEN FRESHWATER PHYTOPLANKTON AND CHYTRID FUNGI (CHYTRIDIOMYCOTA)1

Some chytrids are host‐specific parasiticfungithat may have a considerable impact on phytoplankton dynamics. The phylum Chytridiomycota contains one class, the Chytridiomycetes, and is composed of five different orders. Molecular studies now firmly place the Chytridiomycota within the fungal kingdom. Chytrids are characterized by having zoospores, a motile stage in their life cycle. Zoospores are attracted to the host cell by specific signals. No single physical–chemical factor has been found that fully explains the dynamics of chytrid epidemics in the field. Fungal periodicity was primarily related to host cell density. The absence of aggregated distributions of chytrids on their hosts suggested that their hosts did not vary in their susceptibility to infection. A parasite can only become epidemic when it grows faster than the host. Therefore, it has been suggested that epidemics in phytoplankton populations arise when growth conditions for the host are unfavorable. No support for such a generalization was found, however. Growth of the parasitic fungus Rhizophydium planktonicum Canter emend, parasitic on the diatom Asterionella formosa Hassal, was reduced under stringent nutrient limitation,because production and infectivity of zoospores were affected negatively. A moderate phosphorous or light limitation favored epidemic development, however. Chytrid infections have been shown to affect competition between their algal hosts and in this way altered phytoplankton succession. There is potential for coevolution between Asterionella and the chytrid Zygorhizidium planktonicum Canter based on clear reciprocal fitness costs, absence of overall infective parasite strains, and possibly a genetic basis for host susceptibility and parasite infectivity.     

Observations on a diatom bloom in Loch Leven, Scotland

A study was made on a freshwater planktonic population of centric diatoms, mainly Stephanodiscus rotula (Kütz.) Hendey, in Loch Leven, Kinross, Scotland. The observations were analyzed in relation to changes in water temperature and chemistry, parasitism by a chytridiaceous fungus, grazing by a recently described protozoan and competition from another alga ( Synechococcux sp.). Each of these factors is thought to have affected the recorded rise and fall in diatom population density, in particular, silica limitation, fungal parasitism and protozoan grazing.
The protozoan involved ( Asterocaelum algophilum Canter) forms digestion cysts, the predominantly diatomaceous contents of which suggested very selective feeding but laboratory experiments with dual cultures did not confirm this. They indicated that both the size and quality of algae affected their suitability as food. Feeding activities of Asterocaelum seemed to be more or less confined to a solid surface; grazing rates in cultures shaken to maintain the animals and algae in suspension were low relative to those observed in unshaken cultures. This suggests that much of the grazing occurred on diatoms that had sunk onto the sediments.     

Growth, production and losses of phytoplankton in the lowland River Spree. I. Population dynamics

The seasonal and longitudinal dynamics of dominant phytoplanktonpopulations were observed in the lower Spree (Germany), a lowlandriver draining several lakes. Centric diatoms and cyanobacteriawere the chief components of the total phytoplankton biomassof 16 mg^–1 at the end of the river course (average March-October,1988–89). The cyanobacteria required a retention timeof several weeks in a drained lake to establish stable populations,whereas the other algal populations originated from upstreamimpoundments and slow-flowing river stretches. In spring, thedominant populations increased (the diatoms Nitzschia acicularis,Asterionella formosa, Stephanodiscus hantzschii. Cyclostephanosdubius, Cyclotella radiosa, C.meneghiniana) or kept their abundancesnearly constant (cyanobacterium Limnothrix redekei. diatom Synedraacus), but the biomass of the ‘summer species’ (diatomActinocyclus normanii, cyanobacteria Planktothrix agardhii andAphanizomenonsp.) was halved along the river course. These seasonal differencesin algal growth are probably caused by the changed mixing patternin the regions of origin and thus different pre-adaptation toturbulent mixing. Pronounced dear-water phases occurred regularlyat the end of May. They were intensified along the river coursedespite the lack of mesozooplankton. The dynamics of phytoplanktonwere reflected in the proportions of dissolved and particulatenutrients. The nutrient supply was always high enough to makelimitations of algal growth improbable.     

Prevalence of chytrid parasitism among diatom populations in the lower Columbia River (2009–2013)

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PARASITISM OF ASTERIONELLA FORMOSA HASS. BY A CHYTRID IN TWO LAKES OF THE RAWAH WILD AREA OF COLORADO1

The populations of Asterionella formosa Hass. in two subalpine lakes in the Rawah Wild Area, Colorado, were composed of colonies of five distinct size classes based on frustule length. The spring bloom in these two lakes was dominated by cells of the Beta or Gamma population, whereas the fall pulse was dominated by cells of the Alpha and Beta, or Alpha populations. The Beta population was physiologically unique in that it was parasitized by the chytrid Rhizophidium planktonicum Canter. The development of the parasite population could not be correlated with the concentration or the growth phase of the host population. The occurrrence of parasitism did not noticeiceably affect the population growth patterns of the Asterionella populations.     

PARASITISM OF ASTERIONELLA FORMOSA HASS. BY A CHYTRID IN TWO LAKES OF THE RAWAH WILD AREA OF COLORADO1

The populations of Asterionella formosa Hass. in two subalpine lakes in the Rawah Wild Area, Colorado, were composed of colonies of five distinct size classes Based on frustule length. The spring bloom in these two lakes was dominated by cells of the Beta or Gamma population, whereas the fall pulse was dominated by cells of the Alpha and Beta, or Alpha populations. The Beta population was physiologically unique in that it was parasitized by the chytrid Rhizophidium planktonicum Canter. The development of the parasite population could not be correlated with the concentration or the growth phase of the host population. The occurrence of parasitism did not noticeably affect the population growth patterns of the Asterionella populations.     

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.     

GROWTH RESPONSES OF SEVERAL DIATOM SPECIES ISOLATED FROM VARIOUS ENVIRONMENTS TO TEMPERATURE

Eight diatom species (Chaetoceros pseudocurvisetus Mang ., Stephanodiscus hantzschii Grun ., Skeletonema costatum ( Grev.) Cleve , Asterionella formosa Hass ., Thalassiosira nordenskioeldii Cleve , Detonula confervacea ( Cleve) Gran , Chaetoceros sp., and Nitzschia frigida Grun.) were isolated from various temperature environments ranging from temperate to the Arctic, and their growth responses to temperature were determined. Each species grew over a different temperature range. The lower and upper limits of each species varied from −1.8° to 20° C and from 2° to 30° C, respectively. The width of the growth range of each species. also varied from 3.8° to 25° C, and the growth of these species was observed, as a whole, between a wide temperature range from −1.8° to 30° C .
Within the growth temperature ranges, the growth rate of each species increased with temperature until reaching a maximum, which was followed by a steep decrease up to the upper limit of the growth range. As a result, each species showed a maximum rate at the temperature very near to the upper limit, which was generally higher than the isolation temperature. The specific growth rates were compared among the eight species. The interspecific maximum rate at each temperature exhibited an exponential increase with a Q₁₀ = 1.48. The relative growth rates of each species were calculated by normalizing the specific growth rates with the interspecific maximum rate at each respective temperature. The higher relative growth rates tended to occur at the isolation temperature of each species, suggesting that temperature is a significant control on species distributions in nature .     

Adaptation of the fungal parasite Zygorhizidium planktonicum during 200 generations of growth on homogeneous and heterogeneous populations of its host, the diatom Asterionella formosa

We followed adaptation of the chytrid parasite Zygorhizidium planktonicum during 200 generations of growth on its host, the freshwater diatom Asterionella formosa, in a serial passage experiment. Evolution of parasite fitness was assessed both on a homogenous and heterogeneous host population, consisting of respectively a single new and ten different new host strains. These 10 host strains were genetically different and also varied in their initial susceptibility to the parasite. Parasite fitness increased significantly and rapidly on the new, genetically homogenous host population, but remained unaltered during 200 generations of growth on the heterogeneous host population. Enhanced parasite fitness was the result of faster and more efficient transmission, resulting in higher values of R0 (number of secondary infections). Consequently, parasites that evolved within the uniclonal host population infected significantly more of these hosts than did their ancestors. We thus provide experimental evidence for the widely held view that host genetic diversity restricts evolution of parasites and moderates their harmful effects. Genetically uniform host populations are not only at increased risk from fungal epidemics because they all share the same susceptibility, but also because new parasite strains are able to adapt quickly to new host environments and to improve their fitness.     

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.