Quantifying effects of phytoplankton on the heat budgets of two large limnetic enclosures

1. The aestival heat budgets of two large limnetic enclosures within a small lake in the English Lake District were studied. During summer, these enclosures had different nutrient supplies and consequently different phytoplankton populations. 2. As initial temperature profiles were similar and the incoming surface heat and momentum fluxes for the two enclosures were identical, subsequent changes in the heat budget were assumed to be induced by the biological differences between the enclosures. The proposed mechanism is an increased surface absorption of solar radiation leading to extra surface warming and a consequent excess loss of heat to the atmosphere through long‐wave emittance and sensible and latent heat fluxes, conservatively estimated to be of the order of 10–30 W m^?2. 3. Theoretical calculations show that potential effects on a heat budget could be considerably larger than those observed here. The inherent non‐linearity of the heat fluxes implies that such effects will be more important in warmer lakes than in colder ones. 4. Thermocline depth and strength were also altered by the response to differences in phytoplankton. 5. Any changes in climate or in nutrient loading from the catchment which substantially affect abundance or timing of phytoplankton populations in a lake will consequently also change the thermal structure of the lake.     

Some effects of artificial mixing on the dynamics of phytoplankton populations in large limnetic enclosures

Effects of a series of artificial mixings of part of the watercolumn of a limnetic enclosure (‘Lund Tube’ C, BlelhamTarn, Cumbria) upon the rates of net change in the standingpopulations of selected species of phytoplankton are described.Increase of Sphaerocystis and Anabaena was relatively fasterunder the ‘quiescent’, stratified conditions ofunmixed columns whereas mixing generally favoured the net increaseof diatoms (e.g., Fragilaria) or, at times, of Oscillatoria.The sequences corresponded closely to supposed successionalpathways, which are thus suggested to be naturally regulatedby seasonally-changing physical characteristics of the watercolumn. Performance characteristics of the mixing apparatusand a mathematical expression of the net rate of change in naturalFragilaria populations are appended.     

Experimental manipulations of the phytoplankton periodicity in large limnetic enclosures in Blelham Tarn,English Lake District

This paper reviews the results of experimental manipulations, carried out during the period 1977–1983, on the phytoplankton maintained in the limnetic enclosures at Blelham Tarn, English Lake District. Three categories of manipulations are considered.The effects of variation in the scale and frequency of phosphorus loading (range: 0.3 to 2.5 g P m^–2 a^–1) upon the mean phytoplankton biomass, its seasonal distribution and specific dominance are shown to conform to well-established patterns and relationships observed in natural lakes. Much of the seasonal variability in species dominance occurred independently of nutrient ratios, though carbon availability has been critical at times. Attempts to manipulate the rates of removal of phytoplankton by grazing have confirmed that they act selectively against certain smaller species only, that they alter the rate of successional change, rather than its direction, and that they have little lasting influence upon the total phytoplankton standing crop. Attempts to manipulate rates of sinking loss through artificial enlargement of the epilimnetic circulation also regulated the light-conditions experienced by suspended phytoplankton.Growth-rate relationships to an index of light exposure and to temperature fluctuation are also derived for several species and are related to morphological and physiological characters of the organisms concerned. These interpretations are briefly reviewed in relation to periodic cycles in natural lakes.     

Interrelations between the vertical distribution of Daphnia and chlorophyll a in two large limnetic enclosures

This paper summarises the major factors influencing the daytimevertical distribution of Daphnia hyalina var. laciutris (Sars)in two large experimental enclosures (Lund Tubes). In both tubes,the depth at which the Daphnia aggregated during the day wasclosely related to sub-surface ir-radiance. On a few occasionsaggregations of Daphnia were found in ‘dark’ waterassociated with a deep chlorophyll maximum. On many occasions,however, the animals' light response per se ensured their aggregationat depths of maximum phytoplankton abundance. The most importantfactor influencing the dispersion of animals in the water columnwas the steepness of the light gradient. In turbid water verticalaggregations were well defined, whereas in clear water the animalswere widely dispersed around their depth of maximum abundance.Daphnia also tended to disperse in the water column when theirpopulation density was high or when food was scarce. A simplemodel based on surface irradiance, water turbidity and populationdensity explained the basic pattern of vertical distributionthroughout the season. The implications of these findings arediscussed in relation to the effects of zooplankton grazingand speculations on the adaptive significance of depth regulation.     

Modeling phytoplankton growth rates

Mathematical models of planktonic ecosystems use a variety ofdifferent formulations to relate phytoplankton growth ratesto environmental conditions. Does the formulation influencethe model result? We have modified the model of Fasham, Ducklowarid McKelvie (J. Mar. Res., 34,591–639, 1990) to testhow its results would respond to changes in algal growth rateformulations. The original model uses a Monod relationship betweennutrient concentration and relative growth rate, and a multiplicativerule to combine light and nutrient effects. Use of a Droop formulationfor algal growth rate or a threshold (Blackman's law) mechanismto combine light and nutrient limitation produced significantchanges in simulation results. One important effect was to increasezooplankton population and, as a result, the regenerated production.While there are aesthetic reasons to prefer these alternateformulations, a more accurate formulation will require morelaboratory work on algal physiology. Such laboratory work shouldbe encouraged as an adjunct to modeling work.     

Modeling phytoplankton growth rates

Equation 14 on p. 68 should read: Equation 20 on p. 69 should read:      

Phosphorus recycling in lakes: evidence from large limnetic enclosures for the importance of shallow sediments

• 1 The dynamics of phosphorus in large field enclosures are described from a series of experimental manipulations directed primarily to the study of phytoplankton populations. The enclosures were similar in size but differed in their depth contours. The aim of the present analysis was to detect systematic differences in the phosphorus budgets of deep and shallow systems.
• 2 The sequence of events in each of six year-long experiments is described in terms of algal production. The values are compared with water-column measurements of total phosphorus (TP), the reactive fraction (SRP) and changes in the particulate fraction (PP, assumed to be TP minus SRP). Stoichiometric ratios for larger phytoplankton crops were in the range 85–110 C/P (molar).
• 3 The annual aggregate of new carbon produced in the two uniformly deep enclosures (11.5 m) in relation to the phosphorus supplied was also remarkably close to Redfield stoichiometry, with between 87 and 97mol C yielded per mol P added. However, in the graded enclosure (< 4–13.5 m), the yield was up to twice as great (maximum: 197 mol C mol^-1 P available).
• 4 There are two possible mechanisms. A graded surface permits a greater area of sediment to be in contact with the hypolimnion and for the sediment oxygen demand to deplete a greater hypolimnetic volume; the associated fall in redox would have been adequate to explain the observed increment of dissolved phosphorus to the hypolimnion, associated with the Einsele–Mortimer model (ferric → ferrous iron reduction). There was no clear evidence that this phosphorus supported autotrophic production until late in the year.
• 5 The alternative mechanism is based on the association of phosphorus ‘injection’ to the epilimnion during wind-mixed episodes, when the enhanced shear stress on shallow sediments resuspends fine, superficial deposits and their interstitial waters into the full circulation. Circumstantial evidence suggests this second pathway predominated in the graded enclosure but was ineffective in the uniformly deep enclosures.
• 6 The role of mechanical, aerobic phosphorus cycling in the nutrient economy of lakes, especially small or shallow ones, may have been underestimated in the past but it must be considered when evaluating any proposal to reduce the external nutrient loads to lakes.

     

Sinking losses of phytoplankton in closed limnetic systems

Specific algal recoveries from sediment traps of two differentdesigns and from mud surface deposits of large experimentalenclosures (Lund Tubes) were monitored during 1978 and are analyzedin relation to the vertical and temporal distribution of tendominant phytoplankton populations. Sedimentation accounts fordiffering proportions of the total loss of biomass for differentalgae: between 28 and 100% of diatoms; 15–95% of Eudorina;<4% of populations of small algae (spp. ofAnkyra, Chromulina,Cryptomonas). Rates of diatom loss are also derived from thecomparison of net rates of change (k_n) and the silica uptake-derivedgrowth rate (k¹); intrinsic sinking behaviour may be specificallyregulated in relation to growth conditions. Implications inthe calculation of sedimentary losses and their impact uponthe seasonal periodicity of phytoplankton are briefly discussed.     

Further observations on limnetic zooplankton in a small lake and two enclosures containing fish

SUMMARY. In a small lake (Blelham Tarn, English Lake District) two plastic cylinders, each of which extends from the lake surface to the bottom sediments and encloses 18 000 m³ of water, were used to investigate effects of enclosure on the zooplankton. This paper describes observations from March 1973 to October 1974 and is the second part of a report on a project spanning 4 years.
Seasonal population densities of the more abundant species were essentially similar to those observed in the first part of the project, despite large changes in algal and predator populations. In the enclosures, fish (mostly Perca fluviatilis ) populations are known from observations by divers to have been large in the period under consideration and small in the first part of the project. Populations of Chaoborus larvae in the enclosures were small at all seasons from March 1973 to October 1974.     

Carbon-specific phytoplankton growth rates: a comparison of methods

Measurements of biomass and growth rate of two axenic algalcultures were carried out using three different methodologicalapproaches: the specific ¹⁴C-labelling of chlorophyll a, [³H]adenineincorporation into DNA and net organic carbon assimilation.Time-course experiments revealed that the specific activitiesof chlorophyll a were significantly higher than the specificactivity of total algal carbon in six of seven experiments.When the specific activity of chlorophyll a is used to calculatethe carbon biomass and growth rate, the carbon biomass of thealgae will thus be underestimated and the specific growth ratewill be too high. Determination of growth rates from incorporationof [³H]adenine gave lower values than those obtained from netorganic carbon assimilation and from ¹⁴C incorporation intochlorophyll a. Problems with adenine saturation are suggested.When [³H]adenine is used to measure growth rates in dense algalcultures, additions of >1 µM [³H]adenine are oftenrequired to maximally label the extracellular and intracellularadenine pools and hence DNA.     

In situ growth rates of marine phytoplankton: approaches to measurement, community and species growth rates

Taxonomic structure and biomass weighting are important determinantsof measurable marine phytoplankton community growth potential.Maximal diel-averaged growth rates of communities appear tofall between 3 and 3.6 doublings day^–1. Mean net growthrates are considerably lower. Ranges of community growth ratesmeasured in tropical, sub-tropical and (summer) temperate ecosystemsare similar. There appears to be a broad dichotomy between thegrowth potential of diatom species as compared to non-diatoms.Doubling rates of small diatoms frequently exceed communitybiomass doubling rates by wide margins. Diel-averaged growthrates of large diatoms, microflagellates and non-motile ultraplanktonpopulations are lower and similar in magnitude to communitygrowth estimates. Maximum growth rates of species measured insitu are in good agreement with maximum growth rates measuredin laboratory cultures. High specific productivity of sub-dominantor rare diatom species or assemblages will be diluted in thelower specific growth rates of microflagellate and non-motileultraplankton assemblages. Specific rates of grazing upon speciesand functional groups remain to be quantified, but stabilityof community size and taxonomic structure implies close linkagebetween growth and mortality rates at the species level overtime intervals of several generations.     

Pigment-specific rates of phytoplankton growth and microzooplankton grazing in a subtropical lagoon

Phytoplankton growth and microzooplankton grazing rates wereevaluated in one station in Bahía Concepción,located in the middle region of the Gulf of California, México.We used high-performance liquid chromatography (HPLC) estimationsof phytoplankton pigment signatures to evaluate the annual variationof taxon-specific grazing and growth rates obtained with thedilution technique. Chlorophyll-a (Chl-a) concentrations variedwidely (0.34–3.32 µg L^–1) and showed two maxima,during late spring and autumn, associated with the transitionbetween mixed and stratified conditions. Phytoplankton growthrates varied seasonally with the lowest rates during summer(range: 0.01–2.55 day^–1 for Chl-a; 0.00–3.84day^–1 for Chl-b; 0.26–3.29 day^–1 for fucoxanthin;0.00–6.27 day^–1 for peridinin; 0.00–4.35 day^–1for zeaxanthin). Microzooplankton grazing was an important lossprocess (range: 0.0–1.89 day^–1 for Chl-a; 0.00–3.12day^–1 for Chl-b; 0.26–3.29 day^–1 for fucoxanthin;0.00–2.03 day^–1 for peridinin; 0.00–3.51 day^–1for zeaxanthin). Average grazing rates accounted 68–89%of estimated average phytoplankton pigment-specific growth rates.The analysis of pigment signatures indicates that diatoms anddinoflagellates were the dominant groups, and contrary to expectationfor typical subtropical lagoons, the specific growth rates inBahía Concepción showed a pronounced seasonalvariability, linked to transitional hydrographic conditions.Our results indicate a close coupling between the communitymicrozooplankton grazing and phytoplankton growth rates, withoutselective feeding behavior. These results suggest that microzooplanktonplay a critical role and may significantly modify the availabilityand efficiency of transfer of energy to higher trophic levels.     

Exponential growth rates of phytoplankton calculated from 14C uptake rates: a clarification

Phytoplankton growth rates can be estimated from measurementsof light-dependent [¹⁴C]bicarbonate uptake (photosynthesis)rates using the integrated (logarithmic) form of the exponentialgrowth law. A clanfication of this method is presented herein view of confusions in the recent literature.     

Effect of collection and acclimation period on grazing rates of limnetic zooplankton

Grazing rates of Daphnia sp. and Diaptomus oregonensis measured using an in situ method (Haney 1971) were compared with rates measured by collecting animals in vertical townets and placing them in experimental chambers either immediately or after a 24-h acclimation period. Experiments with acclimation yielded grazing rates of Daphnia that were statistically higher than in situ rates, whereas experiments conducted without acclimation yielded significantly lower rates. In situ grazing rates of Diaptomus were statistically higher than those for both townet techniques; experiments without an acclimation yielded higher rates than those for experiments with a 24-h acclimation period.     

Food-web manipulations influence grazer control of phytoplankton growth rates in Lake Michigan

Stocking piscivorous salmonids in Lake Michigan produced dramaticalterations in food-web structure, including higher numbersof large-bodied zooplankton (especially Daphnia pulicaria),lower summer chlorophyll concentrations and increased watertransparency. Experimental determinations of epilimnetic phytoplanktongrowth rates and of zooplankton grazing rates indicate thatherbivorous zooplankton controlled algal dynamics during thesummer of 1983 because grazers occupied the surface waters throughoutthe day. In 1985, however, both large- and small-bodied Daphniamade approximately equal contributions to total grazer biomass,and all grazers displayed pronounced diel vertical migrations,visiting epilimnetic waters only at night. This prohibited zooplanktonfrom controlling algal dynamics because grazing losses did notexceed phytoplankton growth rates. The changes in zooplanktoncommunity composition and behavior observed in summer 1985 probablyresulted from increased predation by visually orienting planktivorousfish, especially bloater chub (Coregonus hoyi). Effects of food-webmanipulations on phytoplankton dynamics were evident only duringJuly and August. During spring and early summer copepods dominateLake Michigan's zooplankton community. Owing to their smallbody size, copepods are less susceptible to fish predation andexhibit much lower filtering rates than Daphnia. Variabilityin zooplanktivorous fish abundance probably has little effecton phytoplankton dynamics during spring and early summer.     

Comparative carbon-specific ingestion rates of phytoplankton by Acartia tonsa,Centropages velificatus and Eucalanus pileatus grazing on natural phytoplankton assemblages in the plume of the Mississippi River (northern Gulf of Mexico continental shelf)

During four cruises in continental shelf waters of the northern Gulf of Mexico in the winters of 1981–83, we performed quantitative studies on the grazing of the copepods Acartia tonsa, Centropages velificatus, and Eucalanus pileatus, on phytoplankton using natural particulate assemblages as food. Stations were in, or adjacent to the plume of the Mississippi River, thereby providing wide spectra of phytoplankton and suspended riverine particulate concentrations. Phytoplankton cell volume was converted to carbon, and this, coupled with carbon content measurements of these three copepod species, allowed comparisons of daily ingestion effort even though the copepods were of different sizes. Data were expressed in the same units (% of copepod body carbon ingested copepod ^–1 d^–1) for each species. Over similar ranges of phytoplankton carbon concentrations (0.21–92.06 gCl^–1), Acartia tonsa had higher carbon-specific ingestion rates (x = 22.31%, range = 0.08–152.37%) than C. velificatus (x = 2.8%, range = 0.00–31.09 %) or E. pileatus (x = 1.27%, range = 0.10–2.80%). Carbon-specific ingestion rates increased with increasing phytoplankton carbon concentration for A. tonsa (R² = 0.81) and there was no evidence of saturated feeding on the carbon concentrations offered. A similar, but weaker trend was evident for E. pileatus (R² = 0.71), but not C. velificatus (R² = 0.49). Over a wide range of suspended particulate concentrations (10.6–95.2 mg l^–1), there was no systematic effect of particulates on carbon-specific ingestion rate for any of the three copepod species. However, A. tonsa appeared more adept at grazing in highly turbid water than C. velificatus or E. pileatus.     

Measurement of in situ growth rates of phytoplankton under conditions of simulated turbulence

In situ measurement of the growth rates of planktonic populationscan be improved by using dialysis chambers (‘cage cultures’)to avoid shifts in the chemical environment during incubation.Vertical mixing and small-scale turbulence affect the growthof planktonic populations, there fore natural mixing conditionsshould be simulated as closely as possible during the incubation.A new device is described here which combines the advantagesof a dialysis chamber with a programmable vertical mixing regime.Realistic phytoplankton growth rates can thus be measured insitu under con ditions of vertical mixing and small-scale turbulence.The chamber made of transparent, UV-transmitting acrylic glasswas fitted at both ends with permeable polycarbonate membranes.It was moved vertically through the water column by a pocket-sizedlift and rotated simultaneously on its central axis. The methodwas applied to two typos of experiments on growth and lossesof phytoplankton in the River Severn, UK. The first one comparedchanges in biovolume of phytoplankton in a water parcel flowingdownstream (6% h^–1 decline) with those in a simultaneouslyincubated dialysis chamber moved between water surface and riverbottom (7% h^–1 increase). The difference equates to algallosses prevented in the chamber but suffered along the river(mainly sedimentation and grazing of benthic filter feeders).Loss rate of diatoms was three times higher than those of chlorophytes.In another experiment growth of phytoplankton from the mainstream and lateral dead zone was compared under different mixingconditions. Algae from the main stream grew faster than fromthe dead zone. Only cryptophytes preferred calm conditions,all the other algal groups grew faster in chambers moved throughthe water column than in stationary ones. Further possible applicationsin both standing and flowing waters are discussed.     

Taxonomic composition and growth rates of phytoplankton assemblages at the Subtropical Convergence east of New Zealand

Off the eastern coast of New Zealand, warm, saline, nutrient-poorSubtropical Waters (STW) are separated from cool, fresher, relativelynutrient-rich Sub-Antarctic Waters (SAW) by the SubtropicalConvergence (STC). The Chatham Rise, a submarine rise, restrictsthe latitudinal movement of the STC as well as mixing of STWand SAW. Due to this restriction, this sector of the STC ischaracterized by sharp gradients in temperature, macro- (nitrate,silicate and phosphate) and micro- (iron) nutrient concentrations.Shipboard incubations were conducted during austral spring 2000and 2001 to test the hypothesis that these gradients affectthe taxonomic composition and/or growth rates of phytoplanktonon either side of and at the STC. Maximum chlorophyll a concentrationsduring 2000 were 0.39 µg L^–1, but were an orderof magnitude higher in 2001. During both years, STC phytoplanktonwere dominated by diatoms (77% of the total chlorophyll a duringaustral spring 2000 and 70% during spring 2001), whereas cryptophytesand prasinophytes dominated STW assemblages (27 and 36% during2000, and 63 and 17% during 2001). Chlorophyll in the SAW wasdominated by procaryotes and photosynthetic nanoflagellatesduring 2000 (17% procaryotes, 68% nanoflagellates), and by diatomsduring the austral spring 2001 cruise (53%). Growth rates ofthe phytoplankton assemblage were determined by ¹⁴C-labelingof chlorophyll a and photosynthetic pigments. During 2000, temperature-normalizedgrowth rates were near maximal at the STC, and decreased onaverage to less than half of the maximum north and south ofthat front, whereas in 2001 both absolute and relative growthrates were low at all stations. Growth rates did not closelyparallel biomass of the various taxa, suggesting that nutrientlimitation and/or grazing were significantly impacting standingstocks. It appeared that growth was strongly influenced by nutrientsand light, but that biomass was more strongly influenced bygrazing. The STC is a globally important region of enhancedbiomass and productivity; however, the phytoplankton assemblagereflects control by both top–down and bottom–upprocesses that makes a predictive understanding of the area'sbiogeochemical cycles extremely difficult.     

Growth rates of phytoplankton under fluctuating light

• 1 The effect of light fluctuations on the growth rates of four species of freshwater phytoplankton was investigated. Experimental light regimes included constant irradiance and fluctuations of a step function form, with equal proportion of high (maximum of 240 µmol photons m^‐2 s^‐1) and low light (minimum of 5 µmol photons m^‐2 s^‐1) (or dark) in a period. Fluctuations of 1, 8 and 24‐h periods were imposed over several average irradiances (25, 50, 100 and 120 µmol photons m^‐2 s^‐1).
• 2 Growth rate responses to fluctuations were species‐specific and depended on both the average irradiance and the period of fluctuations. Fluctuations at low average irradiances slightly increased growth rate of the diatom Nitzschia sp. and depressed growth of the cyanobacterium Phormidium luridum and the green alga Sphaerocystis schroeteri compared to a constant irradiance.
• 3 Fluctuations at higher average irradiance did not have a significant effect on the growth rates of Nitzschia sp. and Sphaerocystis schroeteri (fluctuations around saturating irradiances) and slightly increased the growth rates of the cyanobacteria Anabaena flos‐aquae and Phormidium luridum (when irradiance fluctuated between limiting and inhibiting levels).
• 4 In general, the effect of fluctuations tended to be greater when irradiance fluctuated between limiting and saturating or inhibiting levels of a species growth‐irradiance curve compared to fluctuations within a single region of the curve.
• 5 The growth rates of species under fluctuating light could not always be predicted from their growth‐irradiance curves obtained under constant irradiance. When fluctuations occur between limiting and saturating or inhibiting irradiances for the alga and when the period of fluctuations is long (greater than 8 h), steady‐state growth‐irradiance curves may be insufficient to predict growth rates adequately. Consequently, additional data on physiological acclimation, such as changes in photosynthetic parameters, may be required for predictions under non‐constant light supply in comparison to constant conditions.

     

A comparison of carbon-specific respiration rates in gelatinous and non-gelatinous zooplankton: A search for general rules in zooplankton metabolism

Using 470 data from the literature the dry weight-specific respiration rates of gelatinous zooplankton (cnidarians, ctenophores and salps) and non-gelatinous zooplankton (mainly crustacea) were converted to carbon-specific values. The resulting carbon-specific respiration rates showed no significant differences between the two groups of zooplankton, indicating similar oxygen requirements per gram of carbon biomass. From this finding, it can be suggested that the differences in the rates of oxygen consumption measured in the two types of zooplankton in the sea can be explained by the carbon biomass ratio between gelatinous and non-gelatinous zooplankton. Furthermore, the low rate of metabolism of gelatinous species compared with that of non-gelatinous animals of the same volume can be attributed predominantly to the relatively low organic matter content in the former. It is recommended that all weight-specific metabolism rates be expressed using carbon as body mass unit (e.g. mg O₂ gC⁻¹ d⁻¹) which enables more accurate comparisons between individuals exhibiting different dry weight/carbon ratios.