Physiological and environmental constraints in the ecology of the planktonic dinoflagellate Ceratium hirundinella

SUMMARY. Constraints upon the vertical distribution of a population of Ceratium hirundinella , in a productive English lake during the summer stratification of 1976, are considered. They are interpreted in relation to vertical gradients of temperature, dissolved oxygen and irradiance, and to rates of photosynthesis and respiration measured as oxygen exchange in long- and short-term exposures. The motile cells tended to aggregate at an intermediate depth in the epilimnion, associated with a relative irradiance level of c . 10% or c . 140 μ einsteins m⁻²s⁻¹ as measured with a horizontal PAR sensor. Higher irradiances, and conditions below the oxycline, were apparently unfavourable, but the intervening layer was severely compressed at the height of summer stratification, when concentrations of inorganic nitrogen were also minimal. Thus the population apparently passed through a critical period, at which the cellular content of chlorophyll-α was much reduced. The preferred irradiance level in the lake corresponded to that at which measured rates of net photosynthesis were maximal. Increased rates of oxygen evolution were measured at higher irradiances in very short exposures; this behaviour may be of importance to cells experiencing vertical movement in nature.     

Some ecological effects of artificial circulation on the phytoplankton

Studies were carried out in Lake Mutek (Mazurian Lakeland) on the effect of artificial aeration and destratification upon quantitative changes in the phytoplankton. These studies were carried out from 1977 until 1983. Two different methods of aeration were used. Low intensity mixing resulted initially in a two-fold, and later on in a four-fold increase of the phytoplankton biomass. Increase of phytoplankton biomass during lake aeration was due to the development of Ceratium hirudinella. Use of a highly effective air-compressor caused an inhibition of algal development, so that biomass dropped to levels noted in the control year. It was found that the effect of aeration depended on the ratio between lake area and effectiveness of the aerator. Only intensive mixing of the water masses resulted in an inhibition of the development of algae. The effect of artificial destratification was also reflected in changes of the species structure, seasonal succession of the algae, and physiological state of the phytoplankton. Artificial circulation stimulated development of algae characterized by relatively high specific weight, i.e. most of all of Pyrrophyta, Bacillariophyceae and some species of Chlorophyta. Various aspects were discussed of the use of direct aeration as a technical method of lake restoration.     

The phased division of the freshwater dinoflagellate Ceratium hirundinella and its use as a method of assessing growth in natural populations

SUMMARY. The nocturnal phasing and partial synchrony of cell division in Ceratium hirundinella was investigated on four occasions for dense planktonic populations in a small productive lake (Esthwaite Water). The population growth rates deduced from the proportion of cells dividing per day are compared with the rates of increase of cell density in the lake. The maximum proportion of Ceratium cells found dividing at any time was 5.8 ± 1.0%, and the time of optimum division was 03.00 hours G.M.T. The daily rate of division during the main phase of population increases was similar to that deduced from the overall population increments at that time, although during the week of collection the increase had apparently ceased temporarily. On the other three occasions, either increase of cell numbers had ceased or the population was declining, but a continued low rate of division ( c . 3% day⁻¹) was found.
The nocturnal division of Ceratium in Esthwaite Water is compared with the division phasing of dinaflagellates described from elsewhere. Some general problems associated with the derivation of estimates of population growth rate from division frequency are also considered.     

Ceratium in Lake Erken: vertical distribution, migration and form variation

External morphological features and growth characteristics of the dinoflagellates Ceratium hirundinella and C. furcoides were investigated in relation to the environmental conditions in Lake Erken. Three distinct population maxima were distinguished, one in June, one in July, and one in September. C. hirundinella dominated the spring and summer populations. C. furcoides was rare in June, but it continuously increased and composed ca. 18% of the Ceratium-cetts in the autumn. The size of a C. hirundinella decreased during the investigation period and a form with a long horn disappeared in June. Ceratium was most numerous during the warmer periods in summer when a distinct thermocline appeared and before the turnover period in September. The alga preferred depths down to 3 m where the blue light dominated and the following parametric magnitudes could be measured: oxygen contents >5 mg 1^-1, pH >7, HCO₃, >1.7 meq. · 1^-1, and a specific conductivity of ca. 250 mS · cm^-1. The algal growth periods displayed doubling times of 2.63 and 4.04 days at surface temperatures of 19°C and 15°C, respectively. The water transparency decreased 3.8 m when Ceratium reached cell densities of 13 individuals · ml^-1 (0–10 m depth). Its vertical migration demonstrated a dial rhythm with accumulation of cells in the surface waters during the day. Complicated patterns of surface avoidance occurred in clear weather and under wind stress.     

Phytoplankton production in relation to physico-chemical conditions in a small, oligotrophic subarctic lake in South Greenland

Phytoplankton production and physico-chemical parameters were measured during the summers of 1976 and 1977 in Lake 95 at 60°52'N, 46°01'W (area 8 ha, z 6.9 m). The duration of lake stratification during summer was weather dependent. Concentrations of total-CO₂ ranged from 2.04 mg l⁻¹ (surface) to 4.02 mg l⁻¹ (bottom during stratification). Nutrient concentrations were generally low, and systematic seasonal variation could not be detected with certainty. Vertical distribution of primary production typically showed a distinct maximum followed by a decline with depth. The low production, both per unit area (max. 60 mg C m⁻² d⁻¹) and per unit volume (max, 14.3 mg C m⁻³ d⁻¹), and the annual production of 4.7 g C m⁻² classified the lake as extremely oligotrophic.     

A seasonal sequence of diel distribution patterns for the planktonic dinoflagellate Ceratium hirundinella in a eutrophic lake

SUMMARY. 1. Diel depth distribution patterns of Ceratium hirundinella were studied during eleven sampling periods, covering the seasonal growth cycle. They were shown to result from short-term periodic or non-periodic external factors, endogenous responses of the alga, and interactions between these.
2. Diel variations in wind stress resulted in the net transport of algae into or out of the sampling position due to lateral water movements. A progressive surface accumulation of Ceratium , leading to a 3-fold increase in cell numbers at the sampling site over a 24 h period, was due to wind-induced upwelling of deeper cells. Near-surface accumulation of Ceratium on a completely overcast afternoon, similar to that associated with migrations, was attributable to advection.
3. Under sufficiently calm conditions, depth-differentiation of Ceratium was regulated by its vertical swimming movements with a diel periodicity. Migration patterns observed in earlier work were confirmed and extended. The alga migrated towards the surface during the daytime and downward during the night; this rhythm had an endogenous component.
4. At high surface illumination, the alga retreated from the surface and formed discrete sub-surface maxima; at low irradiance Ceratium showed positive phototaxis and concentrated near the surface. Under either condition of irradiance Ceratium actively aggregated at depths associated with irradiance levels of about 125–155 μEinsteins m ⁻² s ⁻¹. Downward movement was restricted by anoxic conditions and possibly by thermal/ density gradients. However, given sufficient light penetration, dissolved oxygen and nutrient availability, the thermal density gradient does not apparently eliminate downward movement by Ceratium.     

Phytoplankton ecology in an Antarctic lake

SUMMARY. The ecology of the phytoplankton of Heywood Lake, Signy Island, South Orkney Islands, Antarctica was investigated during 1969–72. The lake, which is ice-covered for 8–10 months per year, is moderately eutrophic due to enrichment by seal excreta.
The annual cycle of the phytoplankton is described. During the winter (approximately May-September), very few algal cells could be detected in the water column and ¹⁴C fixation was below measurable limits. In spring (October-November), a rapidly-growing population of algae caused a large increase in the chlorophyll- a concentration (maximum value 170 mg m⁻²) but carbon fixation remained low, with values <500 mg C m⁻² day⁻¹. The algae contributing to this peak were mainly small chlorophytes and chrysophytes. The summer open-water period (December-March) was characterized by a different phytoplankton population dominated by cryptophytes. Chlorophyll levels were lower ( c . 40 mg m⁻²) but ¹⁴C fixation rates >3 g C m⁻² day⁻¹ were measured on bright days. Values for Assimilation Number were very high (maximum value 10.5 mg C h⁻¹ mg⁻¹ (chlorophyll- a ) in January (1971) though temperatures never exceeded 8°C. In autumn, the phytoplankton regressed to winter levels. Both spring and summer algal populations probably overwinter as resting stages.     

Diel and seasonal physico-chemical fluctuations in a small natural West African lake

SUMMARY. 1. The waters of Opi Lake were very dilute (15–24 μS cm⁻¹at 25°C) and somewhat acid (pH 6.0–6.5), probably a result of the highly leached lateritic soils in its small drainage basin.
2. Annual minimum water temperatures were recorded during the early dry season, not during the rainy season as has been observed in most other West African water bodies. This was attributed to the influence of the Harmattan winds on the lake and the absence of pronounced rainy season flooding, as might occur in reservoirs.
3. A diel cycle of daytime thermal stratification and complete or near-complete night-time destratification occurred throughout the year.
4. During the dry season, when prevailing winds were strong and lake depth shallow, night-time destratification was accompanied by deep circulation. In the rainy season, when prevailing winds were weak and lake depth was greatest, deep vertical circulation apparently did not take place, as evidenced by anoxia in near-bottom waters. This is the reverse of the pattern seen in many tropical lakes.     

The effect of nutrient addition on the phytoplankton community of an oligotrophic lake

This investigation was performed during a 5-yr period (1974–1978) in the oligotrophic Lake Langvatn, Central Norway. In 1975 and 1976 the lake was enriched with a commercial fertilizer, In 1975 increase in phytoplankton biomass was first recorded more than three weeks after the fertilization started, despite a near fivefold increase in the primary production after fertilizer application. The mean seasonal biomass increased from c. 3500 mg wet weight m⁻² in 1974 to 4400 mg in 1975. In 1976 the biomass increased to near 9600 mg ⁻² and the seasonal primary production to 49.0 g C m⁻² (22.2 g C in 1975), despite a reduction in the nutrients added. Chrysophytes constituted the largest share of the seasonal algal biomass in all years, but in fertilization periods cryptophytes dominated in 1975 and diatoms, chlorophytes and cryptophytes in 1976. The highest biomass turnover rate was recorded during a period of cryptophyte dominance. The different biomass and production development in the fertilization years may be explained by a change in the consumer level.     

Population growth rates of two closely related rotifer species: effects of food quantity, particle size, and nutritional quality

SUMMARY. 1. The relationship between population growth rates and the concentrations of several algal species was determined in laboratory experiments with the rotifers Brachionus rubens and B. calyciflorus .
2. The effects of food quantity were well described by a modified Monod model with a threshold for zero population growth. The model parameters depended on particle size and nutritional quality of the food algae. Differences between the rotifer species were significant and reflected their varying food-size preferences.
3. For each rotifer species, thresholds were lowest for algae in the most readily ingested size range. The lowest thresholds were 0.07–0.09 mgC 1⁻¹ with algae of about 5 μm equivalent spherical diameter (ESD) for B. rubens , and 0.19 mgC 1⁻¹ with algae of about 10 μm ESD for B. calyciflorus .
4. Maximal growth rates ( r _max) were slightly below 0.8 day⁻¹ for both rotifers with most algal species. The highest r _max values for both rotifers were observed when Cyclotella meneghiniana was provided as food. With this alga, B. calyciflorus had a significantly higher r_max (1.02 day⁻¹) than B. rubens (0.838 day⁻¹).
5. From a comparison of the relationship between growth rates and ingestion rates, Chlamydomonas reinhardii appeared to be of low nutritional quality for B. rubens .
6. Egg ratios were related to growth rate and were not influenced by the algal food used. Egg development times and average mortality rates were estimated from the relationship between egg ratio and growth rate. B. calyciflorus appeared to have a high average mortality rate (0.383 day⁻¹) compared to B. rubens (0.083 day⁻¹).     

Influence of egg predation and physical disturbance on lake trout Salvelinus namaycush egg mortality and implications for life-history theory

The influence of egg predators and physical disturbance on lake trout Salvelinus namaycush egg mortality was investigated in situ in Lake Michigan where recruitment is below detectable levels and egg predator abundance is high. Comparisons were made with Lake Champlain where recruitment is low and egg predator abundance is also low and with Parry Sound (Lake Huron) where recruitment is moderate and egg predators are in low abundance. A multi-density egg seeding method (100 to 5000 eggs m⁻²) was used to quantify the effect of physical disturbance and egg predation on egg loss. Wind fetch was used as an index of physical disturbance and comparisons across all locations and egg densities suggested that at sites with high wind fetch (>5 km), physical disturbance may be a greater source of egg loss than predation. When analyses were limited to those sites having a wind fetch of <5 km, the percentage of eggs recovered was found to be linearly related to predator density. The strength of this relationship was based largely on egg recovery at 500 and 1000 eggs m⁻² because recovery at lower (100, 250 eggs m⁻²) and very high (5000 eggs m⁻²) densities was not significantly related to predator density. The reason for this is probably that at low egg densities, crayfish Orconectes spp., the major egg predator at most sites, had difficulty finding and consuming eggs and at high egg densities they became satiated. Egg loss was directly related to wind fetch for Lake Michigan and on average six-fold greater than for Parry Sound suggesting that without corresponding changes in fecundity and age structure, lake trout populations in large lakes like Lake Michigan are inherently less productive than those from enclosed inland waters.     

Species composition and biomass of an epipelic algal community in a subarctic lake before and during lake fertilization

The species composition, biomass (measured as algal volumes) and chlorophyll concentration of epipelic algae was studies before (1977) and during (1978–1979) fertilization with phosphorus and nitrogen of Lake Gunillajaure, a small subarctic lake in northern Sweden.
The epipelic biomass, dominated by Cyanophyceae and Bacillariophyceae, was high (5.6–20.1 cm³ m⁻²) at all depths in the lake with the highest values in the hypolimnion (8–13.7 m). Calculated over mean depth it was 20 times higher than that of the phytoplankton. There was no significant increase in biomass during fertilization and neither did the species composition change. The chlorophyll concentration on the other hand were significantly higher in late 1978 and in 1979 which was probably an effect of the declining light climate caused by a large phytoplankton development in the lake. Constant seasonal biomass and species composition indicate a perennial epipelic community in this lake.     

Comparative primary productivity of algal epiphytes on three species of macrophyte in the littoral zone of Lake Ohrid, Yugoslavia

Primary productivity of algal epiphytes on the surfaces of Phragmites, Potamogeton , and Nuphar was measured seasonally from June 1978, through June 1979, in the littoral zone of Lake Ohrid, using ¹⁴C methodology. Surface areas of individual macrophytes were determined throughout the study period through the use of a non-miscible surfactant and a calibration curve of surfactant weight versus known, calculated surface areas.
Mean total surface area available for epiphytic colonization during the study period was 1.032 m² macrophyte surface per m² of littoral zone for Phragmites , 0.810 m² for Potamogeton , and 0.167 m² for Nuphar . Seasonal rates of mean primary productivity of algal epiphytes on Phragmites from the surface to the light-compensation depth ranged from 84–1406 mg C m⁻² littoral zone d⁻¹; ranges for epiphytes on Potamogeton and Nuphar were 77–586 and 69–268 mg C m⁻² littoral zone d⁻¹, respectively. Maximum rates were observed typically during June; minimum rates were observed typically during August to December. Mean daily productivity rates over the 12 month period were for epiphytes on Potamogeton 167.0, on Nuphar 100.4 and on Phragmites 671.2 mg C m⁻² littoral zone d⁻¹. Calculated annual production for epiphytes on Nuphar was 36.65, on Potamogeton 60.95 and on Phragmites 245.0 g C m⁻² littoral zone yr⁻¹. Epiphytic production data were typically considerably higher than production data obtained for littoral and pelagial planktonic algae and compare favorably with published data for epiphytic and periphytic production in Lawrence Lake, Marion Lake, and Borax Lake.     

Abrupt change in primary productivity in a littoral zone of Lake Biwa with the development of a filamentous green-algal community

1. Some characteristics of the photosynthesis and primary production of benthic and planktonic algal communities were investigated in a littoral zone covered with gravel in the north basin of Lake Biwa, paying special attention to the recent development of filamentous green algae (FGA) in the benthic algal community.
2. P_max (maximum gross photosynthesis rate) values of the benthic algal community (0.1–1.2 mg C mg chl. a ⁻¹ h⁻¹) obtained from photosynthesis–irradiance (P–I) curves were lower than those of the planktonic algal community (2.4–11.5 mg C mg chl. a ⁻¹ h⁻¹). This is apparently a result of the high degree of self shading in the benthic algal community and its low turnover as compared with that of the planktonic algal community.
3. Relatively high I_k values (150–200 μmol photon m⁻² s⁻¹) were observed in the benthic algal community only in June–July when a FGA, Spirogyra sp., was abundant. This reflected a photosynthetic characteristic of the Spirogyra itself, in which photosynthesis was saturated at high light intensity.
4. The FGA community established in the layer between planktonic and sessile (benthic algae except for FGA) algal communities. It brought about extraordinarily high organic matter production in the littoral zone at the expense of production in the sessile algal community.     

Laboratory models of diel vertical migration in the dinoflagellate Ceratium hirundinella

SUMMARY. Diel vertical migrations of a dinoflagellate, Ceratium hirundinella , were induced in a laboratory tube (1.63 × 0.15 m) under a light-dark cycle. The timing of vertical migrations differed between cultures in the exponential and stationary phases of growth; the latter showed a greater coincidence with the light regime.
Migration of cells into the surface layers occurred at low values of surface irradiance (<550 μeinsteins m⁻² s⁻¹). At irradiances more closely approaching summer sunshine (> 1300 μE m⁻² s⁻¹) there was a marked avoidance of surface waters, and population maxima were found at depths associated with a relative irradiance level of 10% or c. 150 μE m⁻² s⁻¹). Thermal stratification restricted downward movement of cells into the cooler layers. The combination of high surface irradiance and thermal stratification resulted in large, stable, sub-surface maxima of Ceratium , similar to those observed in natural waters under comparable environmental conditions.     

Divalent cations enhance ammonia excretion in Lahontan cutthroat trout in highly alkaline water

The Lahontan cutthroat trout lives under highly alkaline and saline conditions in Pyramid Lake, Nevada (pH 9.4; 0.2 mmol 1⁻¹ Ca⁺⁺; 7.3 mmol 1⁻¹ Mg⁺⁺). These experiments were conducted to study the possible roles of water Ca⁺⁺ and Mg⁺⁺ concentrations on ammonia excretion in the Lahontan cutthroat trout under highly alkaline conditions. The basic protocol of the experiments was to determine ammonia excretion rates during the following three exposure periods (each of 3-h duration) in sequence: (a) in normal lake water; (b) in soft lake water with the divalent cation concentrations reduced; and (c) in the soft lake water with either Ca⁺⁺ or Mg⁺⁺ (or no divalent cations added) added back at the appropriate lake water concentration. The soft-water exposure caused a significant reduction in ammonia excretion to about half of the control (original lake water) levels. When either Ca⁺⁺ or Mg⁺⁺ was added to the soft water in the third exposure period, the ammonia excretion rates were increased more than twofold back to lake water levels.     

Nutrient retention and transformation in relation to hydraulic flushing rate in a small impoundment

SUMMARY. The fluxes of water, chloride, silicale and various forms of nitrogen and phosphorus into and out of a 3925 ha impoundment (Lake Taiquin) in north Florida were examined over a 3 year period. Annual hydraulic flushing rates for the lake varied by a factor of more than 2 during the study period and provided an opportunity to examine the effect of flushing rate on nutrient cycling within the lake. The results support the hypothesis that nutrient retention and transformation in lakes and impoundments with high flushing rates (> 1 year^-1) are appreciably influenced by annual variations in flushing rates. Higher fractions of input silicate and total phosphorus were retained by Lake Taiquin during the years with average flushing rates (8 year^-1) than during a year with abnormally high flushing rate (16 year^-1). Nitrogen was not appreciably retained during any of the three study years, and thus N: P ratios in the lake outflow were higher than in the inflow.
Phytoplankton productivity was appreciably higher (70%) during the year with the higher flushing rate, apparently in response to higher reactive phosphorus concentrations in the lake and perhaps to a higher standing crop of phytoplankton during that year. Reactive phosphorus and dissolved inorganic nitrogen inputs could account for only about 30% and 25%, respectively, of the phosphorus and nitrogen required to support observed photosynthetic rates in all three study years. Recycling of nutrients apparently accounted for most of the remainder, although nitrogen fixation could not be ruled out as a factor in the nitrogen budget.     

Biomass and annual production of the freshwater mussel Elliptio complanata in an oligotrophic softwater lake

SUMMARY. An extensive survey of Mirror Lake, New Hampshire, was carried out by divers with SCUBA to assess the importance of the freshwater mussel Elliptio complanata in this softwater lake ecosystem. Density (0.032 adults m⁻²), biomass (52 mg m⁻² as dry organic matter) and annual production (6.4 mg m⁻² as dry organic matter) of the mussel population are low when compared with results from other studies, corresponding with the general observation that mussels are scarce in soft, oligotrophic waters. We reject the traditional view that the low mussel density is a result of low calcium concentrations in Mirror Lake, and propose that mussel populations may be regulated by a scarcity of appropriate fish hosts in unproductive lakes. Elliptio complanata is probably not important in the metabolism or biochemistry of the Mirror Lake ecosystem.     

Sediment processing by gizzard shad, Dorosoma cepedianum (Lesueur), in Acton Lake, Ohio, U.S.A.

Gizzard shad are primarily detritivorous in Acton Lake, a 253-ha impoundment in southwestern Ohio, U.S.A. To determine the magnitude of sediment utilization by the gizzard shad population in Acton Lake. I used data on population density and age structure, daily ration, and feeding selectivity in estimating the mass of sediments processed by shad daily from April through November. At densities of 4595–10 645 fish ha⁻¹(wet weight biomass = 90–121 kgha ¹), gizzard shad could process 3.8–23.0 kg of dry sediments ha⁻¹ day ¹. On average throughout the growing season, gizzard shad could process a dry mass of sediments each day equivalent to 13% of shad wet weight biomass. Because of the high rate of sedimentation (> 700 kg dry sediment ha⁻¹ day⁻¹) in Acton Lake, gizzard shad can process < 4% of the freshly deposited sediments each day, and therefore are likely to have little effect on benthic community dynamics in the system.     

The population biology of pike, Esox lucius L., in two gravel pit lakes, with special reference to early life history

The population biology of pike in two gravel pit lakes of contrasting habitat type were studied. The density of pike (1 + and over) in November 1987 in the Main Lake was 7.7 fish ha⁻¹ (19 kg ha⁻¹) and in St Peter's was 71 fish ha'(53 kg ha⁻¹). The age distribution showed that recruitment was consistent in St Peter's but variable in the Main Lake. The main factor causing this difference in recruitment is abundance of aquatic vegetation. Survival up to November 1987 of 0-group pike in the Main Lake was 0.00175% from potential egg deposition, producing 6.6 fish ha⁻¹ (0.83 kg ha⁻¹). In St Peter's survival was 0.00315%, producing 23 fish ha⁻¹ (3.04 kg ha ¹).