Response of Sargasso Sea phytoplankton biomass, growth rates and primary production to seasonally varying physical forcing

The response of phytoplankton biomass, growth rates and primaryproduction to seasonally varying physical forcing was studiedat a station southeast of Bermuda over an 18 month period. Phytoplanktongrowth rates and primary production were measured using thepigment-labeling method, and phytoplankton biomass was calculatedfrom these measurements. Phytoplankton carbon biomass variedsystematically over the year. Highest values were observed duringthe winter and spring. Seasonal variations of chlorophyll (Chi)a in the surface layer could primarily be attributed to variationsin phytoplankton biomass and secondarily to photoacclimation.During the summer period, average values of carbon (C)/Chl ratios(g C g^–1 Chi) ranged from 160 at the surface to 33 atthe 1.6% light level, changes attributed to photoacclimationof the phytoplankton, consistent with the observation that phytoplanktonbiomass did not vary as a function of depth. Phytoplankton growthrates in the surface layer did not vary systematically overthe year, ranging from 0.15 to 0.45 day^–1, in spite ofseasonally varying concentrations of nitrate. Growth rates variedas a function of depth from average values of 0.3 day^–1in the surface layer to <0.1 day¹ at the 1.6% light level.Thus, the primary response of the phytoplankton community tonutrient enrichment during the winter period was an increasein phytoplankton biomass rather than an increase in growth rates.A simple nutrient-phyto-plankton-zooplankton model was usedto explore this phenomenon. The model demonstrated that theobserved response of the phytoplankton to nutrient enrichmentis only possible when phytoplankton growth is not severely limitedby nutrients.     

Bacterial response to seasonal changes in primary production and phytoplankton biomass in Lake Constance

Parameters characterizing bacterial biomass and metabolic activityare compared with phytoplankton biomass and daily primary productionrates throughout the year. Between late March (before the onsetof the phytoplankton spring bloom) and mid-July (diatom maximum),bacterial degradation of organic matter was more closely relatedto phytoplankton productivity than during the rest of the year.Bacterial production (as estimated by amino acid net uptake)was significantly correlated with concentrations of chlorophyll and pheopigments. However, bacterial production was correlatedless closely with primary production and only weakly with bacterialbiomass. Bacterial biomass was also only weakly correlated withprimary production but significantly with pheopigments. Numbersof active bacteria as estimated by autoradiography covariedclosely with bacterial production and cell numbers. Wheneverbacterial production was low, enhanced proportions of aminoacids were respired. Oxygen consumption measurements showedthat the size fraction <3 µm contributed 25–75%to total respiration. On average, bacterial biomass comprised11 % of paniculate organic matter and roughly equalled phytoplanktonbiomass. During the growing season, bacterial production inthe uppermost 20 m comprised about 20% of phytoplankton productionwith large seasonal fluctuations. A tentative carbon budgetof the euphotic zone including primary production, zooplanktongrazing, bacterial production and sedimentation is presented. ¹Present address: Institute of Marine Resources A-018, ScrippsInstitution of Oceanography, University of California, San Diego,La Jolla, CA 92093, USA     

Interannual variability of phytoplankton productivity and related parameters in Lake Constance: no response to decreased phosphorus loading?

In meso-eutrophic Lake Constance (Germany-Austria-Switzerland),phytoplankton bioraass, pigments and water transparency, aswell as primary productivity, have been followed between 1980and 1989. During this period, municipal phosphorus loading declinedsignificantly. Since 1981, soluble reactive phosphorus (SRP)concentrations during deep lake mixing have decreased from 3.0to currently 1 6 mmol m^–3 at a rate of 7% year^–1.Nitrate concentrations, by contrast, continued to rise. Duringthe period of maximum phosphorus loading, flushing through theoutlet and sedimentation were about equally important sinksof phosphorus from the euphotic zone. Recently, however, sedimentationand subsequent burial of P in the bottom deposits contributedabout three-quarters to the overall P-losses from the systemMain reasons for this shift are unchanged settling fluxes ofphosphorus out of the euphotic zone and decreasing concentrationsof total phosphorus in the water. Only during spring, do concentrations of soluble reactive phosphoruswithin the euphotic zone decrease in proportion to the formationof particulate organic matter. Later during the season, euphoticSRP concentrations continue to be low but are no longer matchedby high plankton biomass because phosphorus is efficiently removedby settling of particles In spite of the observed dramatic decreasein phosphorus loading since 1980, chlorophyll concentrationsand water transparency, as well as annual phytoplankton productivity(300 g C m^–2), have not shown a consistent downward trend.However, the intensity of phosphorus regeneration within theeuphoric zone, which can be used as a measure of the degreeof nutrient limitation, is likely to have increased significantlyThe most probable explanation for the insensitivity of importanttrophic state indicators to reduced nutrient loading is that,in Lake Constance, biomass accumulation to a greater extentis controlled by losses, mainly grazing by zooplankton and sedimentation,than by primary resources. This is concluded from the observationthat phytoplankton biomass always falls far short of the nutrient-dependentcarrying capacity of the system.     

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.     

Phytoplankton productivity in turbid waters

Many of the freshwater areas in the world are turbid, due tosuspended inorganic particles. The euphotic depth of the shallowturbid impoundment, Wuras Dam, varies between 0.3–1.3m. This results in a compressed production profile where accuratemeasurements become difficult. Tubes of various lengths havebeen used and usually render higher rates, when compared todiscrete bottle incubations. A tube the depth of the euphoticzone confines the phytoplankton in the light and the rates measuredrepresent the maximal possible under the prevailing conditions.Longer tubes include an aphotic portion and give an idea ofthe magnitude of respiration losses. The depth of the mixinglayer appears to be especially important in turbid systems asthe time spent in the dark, relative to the light is of greatimportance and may be the most important regulating factor insuch waters. ^*This paper is the result of a study made at the Group for AquaticPrimary Productivity (GAP), Second International Workshop heldat the National Oceanographic Institute, Haifa, Israel in April–May1984.     

A model of phytoplankton production in the lower River Rhine verified by observed changes in silicate concentration

The production of phytoplankton in the three main branches andsedimentation areas of the River Rhine in the Netherlands wasanalyzed using a simulation model describing the carbon andsilicate metabolism. This model is based on data derived froma sampling programme in which river water was followed duringdownstream transport. A ‘plug-flow model’ was developed,including sky irradiance and light attenuation in the water,and integrating photosynthetic rates determined in the laboratory.On the basis of the silicate content of diatom-dominated phytoplanktonand silicate regeneration in the river bottom, changes in silicateconcentrations were simulated and found to match observed changesin dissolved silicate. Low sìlicate concentrations wereshown to restrict the maximum population density of diatoms.Depth- and time-integrated rates of photosynthesis were shownto permit multiplication of the phytoplankton at a rate of upto one doubling day^–1 In the primary production periodApril-August 1988. values of 0.48–6.33 g C m^–2 day^–1,close to the few values reported for highly eutrophic riversand lakes, were observed. Model runs, including phytoplanktonproduction and losses, such as respiration, sedimentation andplanktonic grazing, were carried out to simulate the downstreamdevelopment of phytoplankton biomass. These simulations confirmthe view that a substantial part of the phytoplankton biomassand production is grazed or settles in the river delta despiteresidence times of only 52–97 h.     

Phytoplankton Respiration in the Peru Upwelling

Respiratory electron transport system (ETS) measurements weremade on the microplankton in the Peru upwelling system near15°S during March, April, and May of 1977. The close associationbetween chlorophyll- biomass and ETS activity indicate thatthe microplankton were predominantly phytoplankton. Phytoplanktonrespiration average 14% of gross fixed carbon. When zooplanktonrespiration in the euphotic zone is considered, the total planktonrespiration represented an average of 19% of gross primary production. ^*Contribution Number 79002 from the Bigelow Laboratory for OceanSciences.     

Viral lysis modifies seasonal phytoplankton dynamics and carbon flow in the Southern Ocean

Phytoplankton form the base of marine food webs and are a primary means for carbon export in the Southern Ocean, a key area for global pCO₂ drawdown. Viral lysis and grazing have very different effects on microbial community dynamics and carbon export, yet, very little is known about the relative magnitude and ecological impact of viral lysis on natural phytoplankton communities, especially in Antarctic waters. Here, we report on the temporal dynamics and relative importance of viral lysis rates, in comparison to grazing, for Antarctic nano- and pico-sized phytoplankton of varied taxonomy and size over a full productive season. Our results show that viral lysis was a major loss factor throughout the season, responsible for roughly half (58%) of seasonal phytoplankton carbon losses. Viral lysis appeared critically important for explaining temporal dynamics and for obtaining a complete seasonal mass balance of Antarctic phytoplankton. Group-specific responses indicated a negative correlation between grazing and viral losses in Phaeocystis and picoeukaryotes, while for other phytoplankton groups losses were more evenly spread throughout the season. Cryptophyte mortality was dominated by viral lysis, whereas small diatoms were mostly grazed. Larger diatoms dominated algal carbon flow and a single ‘lysis event’ directed >100% of daily carbon production away from higher trophic levels. This study highlights the need to consider viral lysis of key Antarctic phytoplankton for a better understanding of microbial community interactions and more accurate predictions of organic matter flux in this climate-sensitive region.Subject terms: Microbial ecology, Virus-host interactions     

Phytoplankton community growth-rate response to nutrient pulses in a shallow turbid estuary, Galveston Bay, Texas

Phytoplankton growth is a physiological process often limitedby temperature, nutrients or light, while biomass accumulationis a function of growth rates, grazing and deposition. Althoughprimary productivity measurements are usually used to assessresponses to limiting factors, the rates are proportional tobiomass and inversely related to grazing pressure during experimentalincubations. Alternatively, carbon-specific growth-rate determinationsprovide insights into physiological responses without the confoundingeffects of biomass and grazing. The objective of this studywas to quantify the growth-rate responses of phytoplankton toenhanced nutrient availability (nitrate and phosphate) overa range of in situ irradiances. Growth rates were determinedbased on chlorophyll a-specific ¹⁴C-uptake rates by phytoplankton.Phytoplankton demonstrated high (24 h) growth rates when exposedto increased concentrations of limiting nutrients, independentof the surface irradiances (12–41%). Growth-rate responseswere also compared with the biomass (chlorophyll a) responsesand community composition. Observed and estimated phytoplanktonbiomass changes during the incubations differed, emphasizingthe structural role of grazers on the phytoplankton community.The phytoplankton community in Galveston Bay has the potentialto instantaneously respond to nutrient pulses, facilitatingdiatom biomass accumulations in spring and summer and small,flagellated species and cyanobacteria during periods of lownutrient inputs. Thus, Galveston Bay phytoplankton biomass andcommunity composition reflect a dynamic balance between thefrequency of nutrient pulsing and grazing intensity.     

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.     

Pelagic carbon metabolism in a eutrophic lake during a clear-water phase

Dissolved and paniculate organic carbon (DOC and POC, respectively),primary production, bacterial production, bacterial carbon demandand community grazing were measured for 9 weeks in eutrophicFrederiksborg Slotssø. The period covered the declineof the spring bloom, a clear-water phase and a summer phasewith increasing phytoplankton biomass. The process rates andchanges in pools of organic carbon were combined in a carbonbudget for the epilimnion. The POC budget showed a close balancefor both the post-spring bloom and the clear-water phase, whilea surplus was found in the summer phase. Production of POC wasdominated by phytoplankton (2/3) compared to bacteria (1/3)during all phases, and there was a significant correlation betweenphytoplankton and bacterial production rates (r² = 0.48, P <0.039). Bacterial demand for DOC was balanced by productionand changes in the pool of DOC during the decline of the springbloom, but the calculated demand exceeded the supply by 81 and167%, respectively, during the other two periods. The discrepancywas most probably due to an underestimation of bacterial growthefficiency and an overestimation of in situ bacterial productionin carbon units. Production of bacterial substrate by zooplanktonactivity was estimated to be higher than the direct excretionof organic carbon from phytoplankton. The biological successionwas regulated by the balance between area primary productionand community grazing. The clear-water phase was initiated bya combination of low primary production due to low surface irradianceand high community grazing (100 mmol C m^–2 day^–1),which caused a decrease in phytoplankton biomass. However, dueto the high initial phytoplankton biomass, community grazingwas not high enough to cause a significant decrease in areaprimary production. The summer phase was initiated by a decreasein community grazing followed by an increase in phytoplanktonbiomass. Based on these observations and calculations of areaprimary production as a function of chlorophyll concentrations,we suggest that the possibility for zooplankton to regulatephytoplankton biomass in temperate lakes decreases with increasingnutrient level.     

Growth, production and losses of phytoplankton in the lowland River Spree: carbon balance

1. Phytoplankton carbon assimilation and losses (exudation, dark carbon losses) as well as oxygen release and dark community respiration were measured regularly for 2 years at four stations along the lower Spree (Germany). Carbon balance of river phytoplankton was estimated using measured assimilation, metabolic losses and variations in algal carbon along a stretch of river. 2. The light/dark bottle method was modified to simulate vertical mixing. 3. Waxing and waning of phytoplankton populations dominated the load of particulate organic carbon as well as the oxygen budget of the river. 4. Phytoplankton assimilated 310–358 g C m^?2 yr^?1. A mean value of 586 mg C m^?3 day^?1 was fixed in photosynthesis, with 16.7 mg C being exuded during the day and 20.1 mg lost at night. The measured dark respiration was equivalent to only 28% of the daily gross oxygen production of the plankton community. Phytoplankton washed from upstream lakes and reservoirs was not measurably damaged by turbulent transport. 5. In spring, 18–22% of assimilated carbon was used for net biosynthesis of phytoplankton along the river course. At this time, the carbon balance of this part of the Spree was dominated by autochthonous net production. During summer, however, total carbon losses exceeded the intensive carbon assimilation. The decline of algal biomass along the river course in summer was not explicable by measurable physiological losses. The importance of sedimentation and grazing losses is discussed.     

Space-time variability of carbon standing stocks and fixation rates in the Gulf of Maine, along the GNATS transect between Portland, ME, USA, and Yarmouth, Nova Scotia, Canada

The Gulf of Maine North Atlantic Time Series has been run since1998 and is the longest transect time series in the Gulf ofMaine (GoM), USA. Here we use this coastal time series to documentthe space–time variability of hydrography, nutrients,phytoplankton standing stocks and carbon fixation in the GoM,in response to several years of extreme river discharge. Wehypothesize that, during wet years, fresh water input cappedthe surface euphotic layer, impeding the upward diffusion ofnutrients, thus lowering the phytoplankton biomass and carbonfixation rates. Regional algorithms were derived to estimateparticulate organic carbon and carbon fixation. The Howard–Yoderalgorithm was implemented to predict integral primary productionusing satellite ocean color data. Calcification was significantlycorrelated to primary production, thus allowing regional, satellite-derivedcalcification estimates. Total GoM and Georges Bank phytoplanktonphotosynthesis was 38.12 Tg C year^–1 and total calcificationwas 0.55 Tg C year^–1, yielding an overall ratio of calcificationto photosynthesis of 1.44%. Carbon fixation in GoM coastal water(<60 m bottom depth), GoM deep water (>60 m) and GeorgesBank waters (<60 m) averaged 33, 56 and 11% of the totalprimary production of the combined GoM and Georges Bank studyarea, respectively, and 22, 67 and 11% of the total calcificationof the study area, respectively.     

Phytoplankton growth rates in the freshwater tidal reaches of the Schelde estuary (Belgium) estimated using a simple light-limited primary production model

During the course of 1996, phytoplankton was monitored in the turbid, freshwater tidal reaches of the Schelde estuary. Using a simple light-limited primary production model, phytoplankton growth rates were estimated to evaluate whether phytoplankton could attain net positive growth rates and whether growth rates were high enough for a bloom to develop. Two phytoplankton blooms were observed in the freshwater tidal reaches. The first bloom occurred in March and was mainly situated in the most upstream reaches of the freshwater tidal zone, suggesting that it was imported from the tributary river Schelde. The second bloom occurred in July and August. This summer bloom was situated more downstream in the freshwater tidal reaches and appeared to have developed within the estuary. A comparison between phytoplankton growth rates estimated using a simple primary production model and flushing rate of the water indicated that no net increase in phytoplankton biomass was possible in March while phytoplankton could theoretically increase its biomass by 20% per day during summer. Chlorophyllaconcentrations at all times decreased strongly at salinities between 5–10 psu. This decline was ascribed to a combination of salinity stress and light limitation. Phytoplankton biomass and estimated annual net production were much higher in the freshwater tidal zone compared to the brackish reaches of the estuary (salinity > 10 psu) despite mixing depth to euphotic depth ratios being similar. Possible reasons for this high production include high nutrient concentrations, low zooplankton grazing pressure and import of phytoplankton blooms from the tributary rivers.     

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.     

The seasonal productivity of phytoplankton in a hypertrophic, gravel-pit lake

The seasonal time course of phytoplankton primary productivitywas studied weekly in a hypertrophic, gravel-pit lake closeto Madrid, Spain. Chlorophyll a ranged 22–445 mg m^–2.Gross primary productivity attained 0.28±0.14 g C m^–2h^–1 (range: 0.06–0.60), its yearly value being 900g C m^–2, but the shallow euphotic depths and the highplankton respiration ensured that net productivity was generallylow. Respiration losses amounted to 0.31±0.24 g O₂ m^–2h^–1, with phytoplankton respiration roughly attainingone-half of overall plankton respiration. Areal phytoplanktonproductivity and plankton respiration followed a seasonal trendbut this was not the case for photosynthetic capacity. Surfacephotoinhibition was evenly distributed throughout the study.Quantum yields showed an increasing depth trend, but no seasonaltrend. Both P_max and I_k were both temperature- and irradiance-dependent.As compared with lakes of lesser trophic degree, phytoplanktonprimary production in hypertrophic lakes might be increasednot only by higher nutrient contents but also by low chlorophyll-specificattenuation coefficients and low background, non-algal attenuation,thereby allowing for higher areal chlorophyll contents and hencehigher areal productivity. Our study suggests that physical(irradiance and water column stability) as well as chemicalfeatures (dissolved inorganic carbon and soluble reactive phosphorus)may control seasonality of phytoplankton primary productionin this lake despite recent claims that only physical factorsare of significance in hypertrophic lakes. However, this doesnot explain all the variability observed and so a food web controlis also likely to be operating.     

Microplanktonic regeneration of ammonium and dissolved organic nitrogen in the upwelling area of the NW of Spain: relationships with dissolved organic carbon production and phytoplankton size-structure

Ammonium regeneration and dissolved organic nitrogen (DON) releasewere studied experimentally in the euphotic zone of shelf andoceanic waters of NW Spain in relation to coastal upwellingdynamics and the size-structure of phytoplankton communities.Incubations of plankton labelled with [¹⁵N]ammonium were madeduring four cruises, two of which also included size-fractionateddeterminations of chlorophyll a and primary production, andexperimental determinations of production rates of dissolvedorganic carbon (DOC) using ¹⁴C. Inorganic nitrogen concentrationswere mainly related to nitrate enrichment by upwelling pulses,while ammonium concentrations were generally low in all situations.Ammonium did not accumulate in the study area, suggesting adaily time scale coupling between regeneration and uptake. Incontrast, DON largely exceeded inorganic nitrogen in all situationsand generally increased from spring to autumn. Ammonium regenerationwas positively correlated with DON release and both rates showedthe largest variation in summer, with minimum values duringactive upwelling and maximum values when upwelling relaxed.Comparison of DON stocks and rates in different shelf areassuggests that DON release near the coast during summer was morepersistent in the water than DON release in off-shelf and oceanicareas. The carbon:nitrogen ratio of DOC and DON release rateswas highly variable, revealing a large excess of DOC comparedwith DON. This excess can be attributed to either an underestimateof total DON release (as only release from ammonium was measured)or to an enhanced production of carbon-rich organic substancesby diatoms in coastal areas. By considering a broad range oftrophic situations, this study reveals a fundamental differencebetween short term release of DOC and DON by plankton. Physiologicalprocesses (such as carbohydrate exudation by diatoms) seem tobe the cause of large DOC excess, whereas trophic processes(such as grazing) are more likely the cause of enhanced DONrelease.     

AUTECOLOGY OF AN ULTRAPLANKTONIC SHADE ALGA IN LAKE TAHOE2

The ultraplanktonic green alga Monoraphidium contortum Korm. in Lake Tahoe (California-Nevada) demonstrated several ecological and physiological attributes of a genetically adapted shade species. Monoraphidium contortum achieved maximum biomass during deep mixing in winter when light availability was at a minimum. During stratification it was found in maximum abundance in the deep euphotic region, 100–150 m. This species was also distributed through the deep aphotic zone where, despite prolonged darkness, it remained capable of immediate photosynthesis when re-exposed to light levels in the euphotic zone. The spirally twisted cells were grazed by two calanoid copepods in Lake Tahoe as readily as much larger-celled phytoplankton species of less complex morphology. Slow growth rates in combination with high susceptibility to copepod grazing may effectively exclude M. contortum from the upper 75 m, where it was rarely recorded. In culture it showed a marked incapacity to adjust to ‘sun’ conditions but was well adapted to low light regimes. Under a wide range of irradiances, photochemical capacity, photosynthetic capacity and growth rates were low, but cellular pigment content remained high. The ratio of P_max to respiration was at the lower end of the range for shade plants. Genetically distinct sun and shade populations of phytoplankton may play a determining role in major shifts of community structure over depth and time in Lake Tahoe.     

Dynamics of autotrophic picoplankton in Lake Constance

The vertical distribution, biomass concentrations and growthrates of autotrophic picoplankton (APP) were investigated duringthe growing season (March-October) in Lake Constance in differentdepths. Cell numbers determined by epifluorescence microscopyvaried between 1.0 x 10³ and 1.6 times; 10⁵ cells ml^–1depending on season and water depth. Highest concentrationswere recorded above the thermodine in late summer. Numerically,APP consisted almost exclusively of chroococcoid cyanobactena.During lake stratification several peaks of biomass concentrationsoccurred in epilimsietic waters at intervals of 6–8 weeks.In-situ experiments using a dilution technique and dialysisbags revealed that during summer APP population dynamics wereprimarily driven by combined changes of their growth and grazingrates, whereas temperature was less important. Gross growthrates varied between 0.006 and 0.051 h^–1, grazing ratesbetween 0.002 and 0.053 h^–1. On average APP productionwas completely removed by grazing within the microbial community.Ciliates, heterotrophic nanoflagellates and rotifers have beenidentified as the major consumers of APP cells. APP biomassis small compared to larger phytoplankton, ranging from ito5% of total phytoplankton biovolume. Due to its high gross growthrates, which are on the same level as those of free-living pelagicbacteria, APP contributes slightly more to overall primary productionwith maximum percentages of {small tilde}15% in late summer.     

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.