Temporal and vertical variation of chlorophyll a concentration, phytoplankton photosynthetic activity and light attenuation in Lake Kinneret: possibilities and limitations for simulation by remote sensing

The relationship between chlorophyll a (Chl a) and primary productivity(PP) in the uppermost water layer and the water column-based(0–15 m) integral values of those variables were examinedusing measurements taken in Lake Kinneret (Israel) from 1990to 2003. In 81% of all Chl a profiles examined, the distributionwas fairly uniform within the entire 0–15 m water column,and 12.3% of instances showed a prominent subsurface maximum,when the lake phytoplankton was dominated by the dinoflagellatePeridinium gatunense. Chl a can be reliably estimated by remotesensing techniques in the productive and turbid water of LakeKinneret, since Chl a concentration at surface layers can beextrapolated to the entire water column. Light vertical attenuationcoefficient average for wavelengths from 400 to 700 nm, K_d,ranged from 0.203 to 1.954 m^–1 and showed high degreeof temporal variation. The maximal rate of photosynthetic efficiency,P_B^opt [average 3.16 (±1.50)], ranged from 0.25 to 8.85mg C m^–3 h^–1 mg Chl a^–1. Using measured dataof Chl a, P_B^opt, and light as an input, a simple depth-integratedPP model allowed plausible simulation of PP. However, a lackof correlation between photosynthetic activity and temperature(or other variable with remotely sensed potential) renders theuse of models that require input of photosynthetic efficiencyto calculate integrated PP of little value in the case of productiveand turbid Lake Kinneret.     

On the causes of interspecific differences in the growth-irradiance relationship for phytoplankton. II. A general review

The causes of interspecific differences in the µ-l relationshipare examined in the context of a mechanistic model which relatesµ to irradiance in terms of six factors:, k_c photosyntheticquotient (PQ), Chl a:C, respiration and excretion. The effectof cell size on the light saturated growth rate is also considered.It is shown that photosynthetic efficiency and PQ exhibit remarkablylittle interspecific variability, and average 0.024 ±0.005 µg C(µg Chl a)^–1 h^–1 (µEm^–2 s^–1)^–1 and 1.5 ± 0.2 mol 02 molC^–1 (when NO₃^– is the nitrogen source) respectively.Two useful relationships were derived: (i) between growth efficiency,_g and Chl a:C at µ. = 0; (ii) between the compensationintensity, I_c and the Chl a-specific maintenance respirationrate. Both relationships were independent of temperature anddaylength. Species best adapted to growth at low light werefound to exhibit high Chl a:C ratios and low maintenance respirationrates. As a group, diatoms were consistently the best adaptedfor growth at low irradiance. Chiorophytes, haptophytes, chrysophytesand cryptophytes were intermediate in their performance at lowirradiance. Dinoflagellates exhibited extreme diversity, withspecies spanning the spectrum from very good performance atlow irradiance to very poor. A new µ_max-cell carbon relationshipis given based on growth rates normalized to 15°C. Evidenceis presented to show that noise in this relationship can besignificantly reduced by using only carbon-specific growth ratesand using only data for species grown at the same daylength.     

The optical properties of a turbid reservoir and its phytoplankton in relation to photosynthesis and growth (Mount Bold Reservoir, South Australia)

In situ light measurements were used to obtain information oninherent and apparent optical properties. The average verticalattenuation coefficient K_d(ave) varied from 1.1 to 4.6 In unitsm^–1 During three periods the variation in K_d(ave) correlatedwith changes in chlorophyll a concentration and specific attenuationcoefficients K_s, of 0.013, 0.014 and 0.022 m² mg Chl a^–1were calculated. Chlorophyll-specific diffuse absorption coefficients(A,) for these periods were 0.012. 0.013 and 0.017 m² mg Chla^–1 and only varied significantly from estimates of K_sin the period when scattering was intense. Absorption coefficientsa(z^mid) and scattering coefficients b(z^mid) calculated for themid-point of the euphotic zone ranged between 0.45 and 2.9 mand 3.5–52.0 m respectively. Chlorophyll-specific absorptioncoefficients K_a, of 0.005, 0.006 and 0.007 m² mg Chl a^–1and scattering coefficients K_b of 0.05. 0.09 and 0.191 m² mgChl a^–1 were measured during the three periods. The highK_b value occurred when gas-vacuolate cyanobactena were dominant.Algal photosynthesis and light absorption were related throughthe maximum quantum yield _m which varied between 0.019 and 0.11mol C Einstein^–1 while average quantum yields _a, variedbetween 0.006 and 0.024 with a mean of 0.013 mol C Einstein^–1A comparison of changes in the mean irradiance of the mixedzone and chlorophyll concentration indicated that growth waslight limited below 0.04–0.05 Einsteins absorbed mg Chla^–1 day^–1.     

Short-term changes of protozoan control on autotrophic picoplankton in an oligo-mesotrophic lake

In May 1994, we investigated the short-term development of theplanktonic community in the epi- and metalimnion of an oligo-mesotrophiclake (Piburger See, Tyrol), focusing on trophic links betweenprotists and picoplankton, but also including phyto- and zooplankton.Uptake experiments with fluorescently labelled bacteria (FLB)and picocyanobacteria (FIC) were performed in order to comparethe importance of both prey types as carbon sources for bacterivorousprotists. Heterotrophic nanoflagellates (HNF) were responsiblefor {small tilde}90% of total protozoan picoplanktivory (FLB+ FLC); ciliates accounted for {small tilde}10%. A selectivityindex related to prey density showed that both HNF and ciliatesclearly preferred FLC over FLB. The mean cell size of autotrophic(prokaryotic) picoplankton (APP) was nearly three times larger(0.323 µm³) and much less variable than mean bacterialcell volume (0.122 µm³). Although APP biomass was on averageonly 8.6% of total picoplankton biomass, pico-cyanobacteriaaccounted for a mean 15.9% of total HNF carbon uptake. We calculatedthat total HNF grazing could match potential APP maximum growthrates at the beginning of the study period. A strong decreasein HNF individual clearance rate (CR) on APP was clearly relatedto a fall in the percentage of choanofiagellates (from 75 to{small tilde}10% of the HNF community). A simultaneous decreasein HNF biomass and CR was followed by a steep increase in APPabundance; APP abundance and HNF biomass were highly negativelycorrelated both in the epi- and the metalimnion (r_{1 EM} = –0.879,r_{1 META} = =0.907; P = 0.001). Total rotifer abundance increasedby a factor of 50 within 2 weeks and was also negatively correlatedwith HNF biomass (r_{1 EM} = –0.852, P < 0.001; r_{1 META}= –0.659, P < 0.05). HNF grazing was found to exerta strong short-term control on picocyanobacteria and this linkwas probably broken by an increase in metazooplankton, especiallydue to rotifer predation on HNF.     

Growth and development of Neocalanus flemingeri/plumchrus in the northern Gulf of Alaska: validation of the artificial-cohort method in cold waters

In situ growth and development of Neocalanus flemingeri/plumchrusstage C1–C4 copepodites were estimated by both the artificial-cohortand the single-stage incubation methods in March, April andMay of 2001–2005 at 5–6°C. Results from thesetwo methods were comparable and consistent. In the field, C1–C4stage durations ranged from 7 to >100 days, dependent ontemperature and chlorophyll a (Chl a) concentration. Averagestage durations were 12.4–14.1 days, yielding an averageof 56 days to reach C5, but under optimal conditions stage durationswere closer to 10 days, shortening the time to reach C5 (fromC1) to 46 days. Generally, growth rates decreased with increasingstage, ranging from 0.28 day^–1 to close to zero but weretypically between 0.20 and 0.05 day^–1, averaging 0.110± 0.006 day^–1 (mean ± SE) for single-stageand 0.107 ± 0.005 day^–1 (mean ± SE) forartificial-cohort methods. Growth was well described by equationsof Michaelis–Menten form, with maximum growth rates (G_max)of 0.17–0.18 day^–1 and half saturation Chl a concentrations(K_chl) of 0.45–0.46 mg m^–3 for combined C1–3,while G_max dropped to 0.08–0.09 day^–1 but K_chl remainedat 0.38–0.93 mg m^–3 for C4. In this study, in situgrowth of N. flemingeri/plumchrus was frequently food limitedto some degree, particularly during March. A comparison withglobal models of copepod growth rates suggests that these modelsstill require considerable refinement. We suggest that the artificial-cohortmethod is the most practical approach to generating the multispeciesdata required to address these deficiencies.     

On the causes of interspecific differences in the growth-irradiance relationship for phytoplankton. Part I. A comparative study of the growth-irradiance relationship of three marine phytoplankton species: Skeletonema costatum, Olisthodiscus luteus and Gonyaulax tamarensis

Three marine phytoplankton species (Skeletonema costatum, Olisthodiscusluteus andGonyaulax tamarensis) were grown in batch culturesat 15°C and a 14:10 L:D cycle at irradiance levels rangingfrom 5 to 450 µEinst m^–2 s^–1. At each irradiance,during exponential growth, concurrent measurements were madeof cell division, carbon-specific growth rate, photosyntheticperformance (both O₂ and POC production), dark respiration,and cellular composition in terms of C, N and chlorophyll a.The results indicate that the three species were similar withrespect to chemical composition, C:N (atomic) = 6.9 ±0.4, photo-synthetic quotient, 1.43 ± 0.09, and photosyntheticefficiency, 2.3 ±0.1 x 10^–3 µmol O₂ (µgChl a)^–1 h^–1 (µEinst m^–2 s^–1)^–1.Differences in maximum growth rate varied as the –0.24power of cell carbon. Differences in growth efficiency, werebest explained by a power function of Chl a:C at µ = 0.Compensation intensities, ranged from 1.1 µEinst m^–2s^–1 for S. costatum to 35 forG. tamarensis and were foundto be a linear function of the maintenance respiration rate.The results indicate that interspecific differences in the µ–Irelationship can be adequately explained in terms of just threeparameters: cell carbon at maximum growth rate, the C:Chl aratio (at the limit as growth approaches zero) and the respirationrate at zero growth rate. A light-limited algal growth modelbased on these results gave an excellent fit to the experimentalµ–I curves and explained 97% of the observed interspecificvariability. ¹Present address: Lamont-Doherty Geological Observatory Columbiaof University, Palisades, NY 10964, USA     

Planktonic primary production in the German Wadden Sea

By combining weekly data of irradiance, attenuation and chlorophylla concentrations with photosynthesis (P) versus light intensity(E) curve characteristics, the annual cycle of planktonic primaryproduction in the estuarine part of the Northfrisian WaddenSea was computed for a 2 year period. Daily water column particulategross production ranged from 5 to 2200 mg C m^–2 day^–1and showed a seasonal pattern similar to chlorophyll a. Budgetcalculation yielded annual gross particulate primary productionsof 124 and 176 g C m^–2 year^–1 in 1995 and 1996,respectively. Annual amounts of phytoplankton respiration, calculatedaccording to a two-compartment model of Langdon [in Li,W.K.W.and Maestrini,S.Y. (eds), Measurement of Primary Productionfrom the Molecular to the Global Scale. International Councilfor the Exploration of the Sea, Copenhagen, 1993, pp. 20–36],and dissolved production in 1996, were both in the range of24–39 g C m^–2 year^–1. Annual total net productionwas thus very similar to particulate gross production (127 and177 g C m^–2 year^–1 in 1995 and 1996, respectively).Phytoplankton growth was low or even negative in winter. Inspring and summer, production/biomass (Pr/B) ratios varied from0.2 up to 1.7. Phytoplankton growth during the growth seasonalways surpassed average flushing time in the area, thus underliningthe potential of local phytoplankton bloom development in thispart of the Wadden Sea. The chlorophyll-specific maximum photosyntheticrate (P^B_max) ranged from 0.8 to 9.9 mg C mg^–1 Chl h^–1and was strongly correlated with water temperature (r² = 0.67).By contrast, there was no clear seasonal cycle in ^B, which rangedfrom 0.007 to 0.039 mg C mg^–1 Chl h^–1 (µmolphotons m^–2 s^–1)^–1. Its variability was muchless than P^B_max and independent of temperature. The magnitudeand part of the variability of P^B_max and ^B are presumably causedby changes in species composition, as evidenced from the rangeof these parameters found among 10 predominant diatom speciesisolated from the Wadden Sea. The ratio of average light conditionsin the water column (E_av) to the light saturation parameterE_k indicates that primary production in the Wadden Sea regionunder study is predominantly controlled by light limitationand that nutrient limitation was likely to occur for a few hoursper day only during 5 (dissolved inorganic nitrogen) to 10 (PO₄,Si) weeks in the 2 year period investigated.     

Bacterivory in seawater samples estimated by a dual radioactive-labelling technique

A dual radioactive-labelled bacteria technique using Vibrio(DRLV), developed for laboratory studies on bacterivory, hasbeen refined for use at the concentrations of prey and predatorstypcially found at sea. Experiments with estuarine water collectedin spring and in autumn showed that bacterivorous nanoflagellates(HNF) (concentration 1.38±0.35x10³ HNF ml^–1) ingested2.7±0.96 DRLV flagellate1^–1 h^–1 at concentrationsof 0.8–2.2x10⁶ DRLV ml^–1 in the presence of 2.04±0.68x10⁶natural bacteria ml^–1. The method was also applied tosamples collected in October in the Celtic Sea, when on average1 ml of water from the surface layer contained 1.41±0.16x10⁶natural bacteria, 14.6x10³ cyanobacteria, 530±170 HNF,7.3±3.0x103 phototrophic nanoflagellates (1.5–4µm), 49.0±26.5 phototrophic dinoflagellates, 36.3±12.6heterotrophic dinoflagellates and 21.3±9.5 Leucocryptosmarina. Under these conditions the grazing rate in most samplesdid not exceed the coefficient of variation of the method (2%),although we estimate the grazing rate was -1.6 DRLV HNF^–1h^–1 and on one occasion a rate of 2.45 was recorded. Thegross growth efficiency for protein of -30% displayed by naturalHNF means that they could release about     

Photosynthetic Properties of Guard Cell Protoplasts from Vicia faba L.

Guard cell protoplasts were isolated enzymatically from theepidermis of Vicia faba L. and their photosynthetic activitieswere investigated. Time courses of light-induced changes inthe chlorophyll a fluorescence intensity of these protoplastsshowed essentially the same induction kinetics as found formesophyll protoplasts of Vicia. The transient change in thefluorescence intensity was affected by DCMU, an inhibitor ofphotosystem II; by phenylmercuric acetate, an inhibitor of ferredoxinand ferredoxin NADP reductase; and by methyl viologen, an acceptorof photosystem I. Low temperature (77 K) emission spectra ofthe protoplasts had peaks at 684 and 735 nm and a shoulder near695 nm. A high O₂ uptake (175 µmol mg^–1 Chl hr^–1)was observed in guard cell protoplasts kept in darkness, whichwas inhibited by 2 mM KCN or NaN₃ by about 60%. On illumination,this O₂ uptake was partially or completely suppressed, but itssuppression was removed by DCMU, which indicates that oxygenwas evolved (150 µmol mg^–1 Chl hr^–1) photosynthetically.We concluded that both photosystems I and II function in guardcell chloroplasts and that these protoplasts have high respiratoryactivity. (Received January 30, 1982; Accepted May 15, 1982)     

Respiratory ETS activity of plankton in the northwestern Alboran Sea: seasonal variability and relationship with hydrological and biological features

Respiratory electron transport system (ETS) activity was measuredin plankton samples (<200 µm) collected in the NW AlboranSea. Sampling was carried out during seasonal cruises (summerand autumn 2003 and winter and spring 2004) in 12 stations locatedin transects off the coast of Malaga (southern Spain). Thiswork reports for the first time seasonal variations of the Arrheniusactivation energy (Ea) as well as being the first study to addressCO₂ balance in the NW Alboran Sea. These variations were relatedto changes in the phytoplankton community assemblage, whichcould ultimately be caused by the seasonal variability of hydrologicalconditions. ETS activity was significantly higher in summer,coinciding with a higher chlorophyll a (Chl a) concentrationand relatively high levels of particulate organic matter. TheETS:Chl a_total ratios were low during the four seasons, suggestinga high contribution of autotrophic phytoplankton to the respiratoryactivity of planktonic community. Respiratory CO₂ production(RCP) calculated from ETS activity ranged from 4.6 to 28.1 mgC m^–3 day^–1 during the four cruises. Chl a-specificRCP was lower than the maximum photosynthetic rates reportedin the literature for the studied area, suggesting that primaryproduction (PP) and respiration in the water column might beunbalanced.     

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.     

Nitrogen inputs into the euphotic zone by vertically migrating Rhizosolenia mats

Rhizosolenia mats conduct extensive vertical migrations in theoligotrophic central North Pacific (cNP) gyre that permit thesediatoms to acquire nitrate at depth and return to the surfacefor photosynthesis. The ultimate fate of this N within the ecosystemis unknown, but may include remineralization by grazing, lossto depth by sinking biomass, or N excretion by Rhizosoleniamats. Direct release of N by mats into the mixed layer wouldrepresent an upward biological pump that circumvents the diffusionbarriers and nutrient sinks at the base of the oceanic euphoticzone. We examined Rhizosolenia mat N release along a transect(28–31° N) in the summer of 2002 (Hawaii to California)and 2003 (Hawaii to west of Midway Island) using sensitive fluorometricand chemiluminescence methods. Nitrate, NO₂^– and NH₄⁺release was determined. Nitrate and NH₄⁺ release by the matsoccurred in both 2002 (22.84 ± 6.04 and 3.69 ±1.74 nmol N µg^–1 Chl a h^–1, respectively)and 2003 (23.74 ± 3.54 and 3.60 ± 0.74 nmol Nµg^–1 Chl a h^–1, respectively). Nitrite releaseonly occurred in the 2003 summer period but occurred in bothyears when Fe chelators were added. Fv/Fm values decreased westwardin 2003 suggesting a gradient of increasing physiological stresstowards the west. The various physiological measures are consistentwith concurrent Fe stress; however, other possibilities exist.Nitrate excretion was the dominant form of N release in bothyears and provided a substantial addition to the ambient nitratepool in the mixed layer. Rhizosolenia mat nitrate release suppliesat least 4–7% of the nitrate pool on daily basis, andpossibly as much as 27%. Rhizosolenia mats are part of a largephytoplankton community that appears to migrate, and rates couldbe significantly higher. Literature reports suggest little orno nitrification in the upper euphotic zone, and thus biologicaltransport and release of nitrate may be a major source to thisregion. This N release is uncoupled from upward CO₂ transportand, like N₂ fixation, provides a component of the N pool availablefor net carbon removal.     

Spatial distribution of transparent exopolymer particles in the Bransfield Strait, Antarctica

Transparent exopolymer particles (TEP) are recognized to playan important role in the flux of exported carbon to the deepocean. However, there is little information on how TEP standingstocks are affected by different hydrographic conditions andother relevant ecological factors in situ. This lack of knowledgeis particularly serious for the Southern Ocean. During Australsummer 1999, the Strait of Bransfield presented high mesoscalevariability. Two fronts were present, the Bransfield hydrographicfront and a slope front along the South Shetland Islands andseveral mesoscale anticyclonic eddies and/or frontal meanders.The spatial distributions of biological properties were largelyaffected by this complex hydrography. Chlorophyll a (Chl a)(0.05–4.81 µg L^–1), TEP (from undetectableto 346 µg GXeq L^–1) and heterotrophic bacteria (HB)(1.7–9.4 x 10⁵ cells mL^–1) were positively correlateddespite the wide hydrographic heterogeneity of the BransfieldStrait. Higher abundances of autotrophic biomass, and correspondlyhigher TEP and heterotrophic bacteria (HB), were found in themore stratified waters. TEP spatial distribution was mostlyrelated to the abundance of autotrophic biomass although localhigh TEP concentrations were not matched by similarly high valuesof Chl a in some areas where diatoms were relatively abundant.     

Seasonal and interannual variability of chlorophyll a and primary production in the Equatorial Atlantic: in situ and remote sensing observations

The seasonal variability of phytoplankton in the EquatorialAtlantic was analysed using Sea-viewing Wide Field-of-view Sensor(SeaWiFS)-derived chlorophyll a (Chl a) concentration data from1998 to 2001, together with in situ Chl a and primary productiondata obtained during seven cruises carried out between 1995and 2000. Monthly averaged SeaWiFS Chl a distributions werein agreement with previous observations in the Equatorial Atlantic,showing marked differences between 10° W in the EasternTropical Atlantic (ETRA) and 25° W in the Western TropicalAtlantic (WTRA) provinces (Longhurst et al. 1995. J. PlanktonRes., 17, 1245–1271). The seasonal cycle of SeaWiFS-derivedChl a concentration calculated for 0–10° S, 0–20°W (ETRA) is consistent with in situ Chl a measurements, withvalues ranging from 0.16 mg m^–3, from February to April,to 0.52 mg m^–3 in August. Lower variability was observedin 10° N–10° S, 20–30° W (WTRA) whereminimum and maximum concentrations occurred in April (0.15 mgm^–3) and in August (0.24 mg m^–3), respectively.A significant empirical relationship between depth-integratedprimary production and in situ measured sea surface Chl a wasfound for ETRA, allowing us to estimate the seasonal cycle ofdepth-integrated primary production from SeaWiFS-derived Chla. As for Chl a, this model was verified in a small area ofthe Eastern Equatorial Atlantic (0–10° S, 0–20°W), although in this instance it was not completely able todescribe the magnitude and temporal variability of in situ primaryproduction measurements. The annual euphotic depth-integratedprimary production rate estimated for ETRA by our empiricalmodel was 1.4 Gt C year^–1, which represents 16% of theopen ocean primary production estimated for the whole AtlanticOcean.     

Size-fractionated primary production studies in the vicinity of the Subtropical Front and an adjacent warm-core eddy south of Africa in austral winter

Results are presented of size-fractionated primary productionstudies conducted in the vicinity of the Subtropical Front (STF),an adjacent warm-core eddy, and in Sub-antarctic waters duringthe third South African Antarctic Marine Ecosystem Study (SAAMESIII) in austral winter (June/July) 1993. Throughout the investigation,total chlorophyll (Chl a) biomass and production were dominatedby small nano- and picophytoplankton. No distinct patterns intotal Chl a were evident. At stations (n = 7) occupied in thevicinity of the STF, total integrated biomass values rangedfrom 31 to 53 mg Chl a m^–2. In the vicinity of the eddy,integrated biomass at the eddy edge (n = 3) ranged from 24 to54 mg Chl a m^–2 and from 32 to 43 mg Chl a m^–2 inthe eddy (n = 2). At the station occupied in the Sub-antarcticwaters, total integrated biomass was 43 mg Chl a m^–2.Total daily integrated production was highest at stations occupiedin the vicinity of the STF and at the eddy edge. Here, totalintegrated production ranged from 150 to 423 mg C m^–2day^–1 and from 244 to 326mg C m^–2 day^–1, respectively.In the eddy centre, total integrated production varied between134 and 156 mg C m^–2 day^–1. At the station occupiedin the Sub-antarctic waters, the lowest integrated production(141 mg C m^–2 day^–1) during the entire survey wasrecorded. Availability of macronutrients did not appear to limittotal production. However, the low silicate concentrations duringthe survey may account for the predominance of small nano- andpicophytoplankton. Differences in production rates between theeddy edge and eddy core were related to water column stability.In contrast, at stations occupied in the vicinity of the STF,the control of phytoplankton production appears to be relatedto several processes, including water column stability and,possibly, iron availability.     

Microzooplankton grazing in a coastal embayment off Concepcion, Chile, (~36{degrees} S) during non-upwelling conditions

The impact of grazing by natural assemblages of microzooplanktonwas estimated in an upwelling area (Concepción, Chile)during the non-upwelling season in 2003 and 2004. Seawater dilutionexperiments using chlorophyll a (Chl a) as a tracer were usedto estimate daily rates of phytoplankton growth and microzooplanktongrazing. Initial Chl a concentrations ranged from 0.4 to 1.4mg Chl a m^–3 and phytoplankton prey biomass and abundancewere numerically dominated by components <20 µm. Phytoplanktongrowth and microzooplankton grazing rates were 0.19–0.25day^–1 and 0.26–0.52 day ^–1, respectively.These results suggest that microzooplankton exert a significantremoval of primary production (>100%) during the non-upwellingperiod.     

Indirectly fluorescently labelled flagellates (IFLF): a tool to estimate the predation on free-living heterotrophic flagellates

A procedure was developed to estimate the direct grazing impacton free-living heterotrophic nanoflagellates (HNF). Culturedflagellates were labelled by feeding on brightly fluorescingbacteria (FLB) and then offered as indirectly fluorescentlylabelled flagellates (IFLF) to potential predators of HNF. Thenumber of FLB in the predators' food vacuoles could be convertedinto IFLF uptake and consumption of HNF. This new techniquewas used to study the HNF-ciliate relationship in the pelagiczone of Lake Constance. Three groups of ciliates were detectedas HNF grazers: small representatives of the genus Strobilidium.a small Haltena-like ciliate (probably Halteria grandinella)and a Codonella sp. Tintinnidium sp. group The ingestion ofHNF by these groups of ciliates ranged between 3 and 15, 3 and39, and 3 and 7 HNF ciliate^–1 h^–1; respectively.The IFLF method allows the direct determination of ingestedflagellate prey in the food vacuoles of their predators. Becauseindigenous living prey organisms were used, tracer discriminationcan be reduced.     

Time-Resolved Spectroscopy of Chlorophyll-a like Electron Acceptor in the Reaction Center Complex of the Green Sulfur Bacterium Chlorobium tepidum

The absorption changes of chlorophyll (Chl) a-like pigments(C670) were studied by ns-ms laser spectroscopy at 77 K in theuntreated and urea-treated homodimeric reaction center (RC)complex of the green sulfur bacterium Chlorobium tepidum. Theuntreated RC complex contained 9 molecules of C670 in additionto 41 molecules of Bchl a and 0.9 molecules of menaquinone-7per one primary electron donor Bchl a dimer (P840). Upon photo-oxidationof P840, C670 showed an absorption change of a red-shift withan isosbestic wavelength at 668 nm. The absorption change ofP840 decayed with time constants (t_1/e) of 55 and 37 ms at 283and 77 K, respectively, and was assigned to represent the chargerecombination between P840⁺ and FeS^–. In the urea-treatedRC complex, a bleach peaking at 670 nm with a shoulder peakat 662 nm, which is ascribable to the reduced primary electronacceptor A₀^–, was detected after the laser excitationin addition to the shift at 668 nm indicating the formationof the P840⁺A₀^– state. The P840⁺A₀^– state decayedwith a t_1/e of 43 ns at 77 K and produced a triplet state p840^Tdue to the suppression of the forward electron transfer. Theseresults indicate the two different types of C670 species inthe RC complex; the one peaking at 670 nm functions as A⁰, whilethe other peaking at 668 nm shows the electrochromic shift,which presumably functions as the accessory pigment locatedin the close vicinity of P840. (Received May 17, 1999; Accepted July 14, 1999)     

Comparison of east and west chlorophyll a standing stock and oceanic habitat along the Transition Domain of the North Pacific

Chlorophyll (Chl) a was measured every 10 m from 0 to 150 min the Transition Domain (TD), located between 37 and 45°N,and from 160°E to 160°W, in May and June (Leg 1) andin June and July (Leg 2), 1993–96. Total Chl a standingstocks integrated from 0 to 150 m were mostly within the rangeof 20 and 50 mg m^–2. High standing stocks (>50 mg m^–2)were generally observed westof 180°, with the exceptionof the sporadic high values at the easternmost station. Thetotal Chl a standing stock tended to be higher in the westernTD (160°E–172°30'E) than in the central (175°E–175°W)and eastern (170°W–160°W) TD on Leg 1, but thesame result was not observed on Leg 2. It was likely that largephytoplankton (2–10 and >10 µm fractions) contributedto the high total Chl a standing stock. We suggest that thehigh total Chl a standing stock on Leg 1, in late spring andearly summer, reflects the contribution of the spring bloomin the subarctic region of the northwestern Pacific Ocean. Thedistribution of total Chl a standing stock on Leg 2 was scarcelyaffected by the spring phytoplankton bloom, suggesting thattotal Chl a standing stock is basically nearly uniform in theTD in spring and summer. Moreover, year-to-year variation inthe total Chl a standing stock was observed in the western TDon Leg 1, suggesting that phytoplankton productivity and/orthe timing of the main period of the bloom exhibits interannualvariations.     

Bulk-phase viscoelastic properties of seawater relationship with plankton components

The viscous and elastic moduli at different shear rates, togetherwith various biological oceanographic properties, were determinedin seawater from different hydrological layers in the southernNorth Sea in June. The biological oceanographic parameters includedPhaeocystis and Noctiluca abundances, chlorophyll a level (Chl),bacteria. HNAN and aggregate volume fraction. The plankton wasjointly dominated by Phaeocyslis sp. and Noctiluca scinullans.Noctiluca abundance showed no correlation with any other biologicalor viscoelastic parameter, but Phaeocystis abundance correlatedstrongly. The other biological parameters correlated with Phaeocystisand with each other positively and mostly significantly. Overall,viscoelasticity correlated more strongly with Chl than withany other biological parameter. For non-microlayer samples,the excess complex (viscoelastic) modulus (µ.Pa) G^*_E =2.0 x Ch1^1–3 (Chl in mg m^–3). Viscous and elasticmoduli also correlated closely with each other. For a givenvalue of Chl. the microlayer samples were 6.5 or 14 times (dependingon the estimation method) more viscoelastic than in bulk-phasesamples. Viscoelasticity in samples of settled benthic ‘fluff’were lower even than bulk-phase samples, but this differencewas not significant. Comparison with Mediterranean data on viscoelasticity(Jenkinson. Oceanol. Acta, 16, 317–334, 1993), using publishedvalues for phytoplankton biomass (Wiadnyana, J. Rech. Océanogr.,17, 1–6, 1992), suggests that the relationship betweenChl (or phytoplankton biomass) and viscoelasticity might begeneral. This apparent biomodification of the viscosity andelasticity of seawater is discussed in relation to its likelyimpact on turbulence and plankton ecology.