Summer dynamics of the deep chlorophyll maximum in Lake Tahoe

Vertical profiles of chlorophyll and phytoplankton biomass weremeasured in Lake Tahoe from July 1976 through April 1977. Adeep chlorophyll maximum (DCM) persisted during summer and earlyautumn (July—October) near 100 m, well below the mixedlayer and at the upper surface of the nitracline. The DCM coincidedwith the phytoplankton biomass maximum as determined from cellcounts. In addition, the composition of the phytoplankton assemblagewas highly differentiated with respect to depth. Cyclotellastelligera was the predominant species in the mixed layer whilethe major species in the DCM layer included C. ocellata andseveral green ultraplanktonic species. In situ cell growth playsa substantial role in maintaining the DCM, but sinking of cellsfrom shallower depths and zooplankton grazing above the DCMmay contribute to the maintenance of the DCM. Calculations supportthe interpretation that the summer DCM persists at the boundarybetween an upper, nutrient-limited phytoplankton assemblageand a deeper, light-limited assemblage.     

Does the chlorophyll a content of phytoplankton vary with trophic status in lakes on the New Zealand central volcanic plateau?

Recently, it has been shown that ratios of chlorophyll a toparticulate phosphorus (Chl a/PP) and chlorophyll a to particulatenitrogen (Chl a/PN) were significantly higher in eutrophic thanoligo/mesotrophic waters in 17 lakes on the central volcanicplateau, North Island, New Zealand. This difference was thoughtto be due to an increase in the chlorophyll a content of phytoplanktonin these eutrophic lakes. Corresponding measurements of chlorophylla and phytoplankton cell volume made during this study do notsupport this hypothesis. However, ratios of chlorophyll a toadenosine triphosphate and estimates of percentage phytoplanktonbiomass were significantly higher (P<0.05) in our eutrophicthan oligo/mesotrophic samples, suggesting that Chl a/PP andChl a/PN may be high in eutrophic waters simply because phytoplanktoncomprise more of the total microbial biomass. This hypothesisis supported by a strong linear relationship (r=0.88, P<0.001)between Chl a/PP and percentage phytoplankton biomass in sixof our study lakes where corresponding measurements were made.     

Factors affecting phytoplankton production over the Campbell Plateau, New Zealand

The Campbell Plateau, which covers approximately 600,000 km²at depths ranging from 0–500 m on the rises to 1500 mat the plateau edge, possibly encompasses a unique combinationof bathymetric and hydrological features. Nutrients are in good supply and do not limit primary production.Over areas deeper than 450 m chlorophyll a concentrations aregenerally low but are greater where the stability of the surfacewaters is greater. Chlorophyll a reaches maximum concentrationsin areas shallower than 500 m (e.g., Pukaki Rise and BountyPlateau). Water over the Campbell Plateau generally has low stability,the result presumably of bathymetrically induced mixing andthe turbulent meteorological regime. Phytoplankton-poor wateris apparently introduced into this area where water column stabilityis insufficient to support rapid phytoplankton growth. Shallowbathymetric features appear to confine the phytoplankton nearthe surface and enhance phytoplankton production. It is postulated that current speed and direction is responsiblefor the skewness of the relationship between bottom depth andthe distribution of chlorophyll a and differences between eachbathymetric feature.     

Estimating chlorophyll a abundance from the 'phytoplankton colour' recorded by the Continuous Plankton Recorder survey: validation with simultaneous fluorometry

The green colour (measured with reference to standard colourcharts) of sections of the Continuous Plankton Recorder (CPR)filtering silk was compared with estimates of chlorophyll aconcentration derived from a fluorometer mounted on the CPRduring seven tows in the North Sea between February and May1991. After the green colour was assessed, the abundance ofphytoplankton cells on the filtering silks was quantified bymicroscope analysis. Data were collected for 115 10-nautical-milesamples over a total of seven cruises. For these 115 samples,there was only a weak (F_1.113 = 3.8, P = 0.05, r² = 0.03) positiverelationship between the colour of the filtering silk and thechlorophyll a concentration. However, when this comparison wasrestricted to four tows (68 10-nautical-mile samples) wherethe recorded phytoplankton cell abundance on the silks was verylow, there was a highly significant (F_1.66 /,69.1, P < 0.001,r² = 0.51) positive relationship between the silk colour andthe chlorophyll a concentration. By measuring the relative colourintensity of CPR standard colour categories and quantifyingthe individual variation in the assessment of colour, a theoreticalmodel was developed which pedicted that if the silks were colouredin direct proportion to the chlorophyll a concentration in thewater, then the expert r² for the relationship between silkcolour and chlorophyll a concentration would be 0.62. The greencolour recorded by the CPR survey was therefore identified asa quantitative index of chlorophyll a concentration, but onlywhen numbers of phytoplankton cells on the CPR silks are nothigh.     

Biomass-pigment relationships in potamoplankton

During most of the growing season of 1994, pigment content,as determined by HPLC analysis of algal sample extracts, wasfollowed in the River Meuse (Belgium) potamoplankton. The concentrationof some algal pigments (chlorophylls a and b, fucoxanthin, lutein,echinenone and alloxanthin) was related to biomass estimatesof total phytoplankton and of major taxonomic components (diatoms,green algae, cyanobacteria and cryptomonads). Highly significantlinear regressions were obtained for chlorophyll a-total biomass,fucoxanthin-diatoms, lutein-green algae, chlorophyll b-greenalgae. However, no relationship was found for cyanobacteriaor cryptomonads and their specific pigments, which may be attributedto poor accuracy of biomass estimates for these non-dominantalgae. In conclusion, the good relationship found for dominantalgae and their specific pigments confirms the value of pigmentsas quantitative markers of phytoplankton, as detected in othermarine and freshwater environments.     

Size and species-specific primary productivity and community structure of phytoplankton in Tokyo Bay

Combined methods of size fractionation and single-cell isolationwere used to investigate the seasonal variation of phytoplanktondynamics in Tokyo Bay with an emphasis on primary productivity.Red tides occurred in Tokyo Bay from spring to autumn; a diatom,Skeletonema costatum, and a raphidophycean, Heterosigma akashiwo,were the most important primary producers. Small diatoms andflagellates, including these species, were dominant and showedrapid changes of phytoplankton community structure within severaldays in summer. The nanoplankton (3–20 µm) fractioncontributed most to chlorophyll a concentration and primaryproductivity during spring to autumn, whereas the microplankton(>20 µm) contribution was remarkable in winter. Picoplankton(<3 µm phytoplankton) remained relatively constantthroughout the year. A significant reverse relationship wasobtained between assimilation rate and chlorophyll a contentfor the total and nanoplankton population; the assimilationrate was high at the initial phase of the bloom, then decreasedto a minimum level at the peak of the bloom. Factors controllingthe reduction of assimilation rates at the peak, and changesin phytoplankton community structure, are discussed.     

Response of phytoplankton and bacteria to nutrients and zooplankton: a mesocosm experiment

Although both nutrient inputs and zooplankton grazing are importantto phytoplankton and bacteria in lakes, controversy surroundsthe relative importance of grazing pressure for these two groupsof organisms. For phytoplankton, the controversy revolves aroundwhether zooplankton grazers, especially large cladocerans likeDaphnia, can effectively reduce phytoplankton populations regardlessof nutrient conditions. For bacteria, little is known aboutthe balance between possible direct and indirect effects ofboth nutrients and zooplankton grazing. However, there is evidencethat bacteria may affect phytoplankton responses to nutrientsor zooplankton grazing through direct or apparent competition.We performed a mesocosm experiment to evaluate the relativeimportance of the effects of nutrients and zooplankton grazingfor phytoplankton and bacteria, and to determine whether bacteriamediate phytoplankton responses to these factors. The factorialdesign crossed two zooplankton treatments (unsieved and sieved)with four nutrient treatments (0, 0.5, 1.0 and 2.0 µgphosphorus (P) l^–1 day^–1 together with nitrogen(N) at a N:P ratio of 20:1 by weight). Weekly sieving with 300µm mesh reduced the average size of crustacean zooplanktonin the mesocosms, decreased the numbers and biomass of Daphnia,and increased the biomass of adult copepods. Nutrient enrichmentcaused significant increases in phytoplankton chlorophyll a(4–5x), bacterial abundance and production (1.3x and 1.6x,respectively), Daphnia (3x) and total zooplankton biomass (2x).Although both total phytoplankton chlorophyll a and chlorophylla in the <35 µm size fraction were significantly lowerin unsieved mesocosms than in sieved mesocosms, sieving hadno significant effect on bacterial abundance or production.There was no statistical interaction between nutrient and zooplanktontreatments for total phytoplankton biomass or bacterial abundance,although there were marginally significant interactions forphytoplankton biomass <35 µm and bacterial production.Our results do not support the hypothesis that large cladoceransbecome less effective grazers with enrichment; rather, the differencebetween phytoplankton biomass in sieved versus unsieved zooplanktontreatments increased across the gradient of nutrient additions.Furthermore, there was no evidence that bacteria buffered phytoplanktonresponses to enrichment by either sequestering P or affectingthe growth of zooplankton.     

Spatio-temporal dynamics of chlorophyll in the open Baltic Sea

Five charts of the chlorophyll and hydrographic fields by verticalprofiles of in situ fluorescence and CTD were made on a stationarygrid of 20 ? 25 nautical miles with a 5-mile spacing in theopen Baltic Sea. Both chlorophyll levels and variability weremaximal close to the spring bloom. High chlorophyll levels insummer are sustained by recurrent nutrient injections from thedeep saline layer. Two of the surveys showed close couplingbetween the coarse-scale (10 km) chlorophyll distribution andthe hydrographic structure determining the intensity of nutrienttransfer. Vigorous advection, stirring and current shear, associatedwith a strong mesoscale eddy, probably dominated the chlorophyllpattern on three surveys. The upward velocities in the cycloniceddy resulted in accumulation of phytoplankton in the aphoticzone. Intensive heat input from the surface caused a suddensinking of the phytoplankton and the formation of a pronouncedsub-surface chlorophyll maximum.     

HPLC analysis of phytoplankton pigments from Lake Kinneret with special reference to the bloom-forming dinoflagellate Peridinium gatunense (Dinophyceae) and chlorophyll degradatio products

Photosynthetic pigments extracted from the paniculate materialof the water column of Lake Kinneret were studied throughoutthe periods of May 1988-June 1989, and November 1993-November1994, by means of HPLC. The temporal and vertical variationof the pigment suite found agreed with the microscopically determinedphytoplankton record. The regression calculations of taxon-specificbiomass with the corresponding signature pigments suggest thatpigment analysis may be a useful tool for the monitoring ofbloom-forming species, e.g. the dinoflagellate Peridinium gatunenseNygaard. The HPLC pigment analysis permitted the identificationand quantification of chlorophyll degradation products, providingfor the first time information about their composition in LakeKinneret. Chlorophyllide a was the major detectable degradationproduct of chlorophyll a, varying between 1 and 9% of the chlorophylla concentration. Other chlorophyll a derivatives appeared mostlyin minor quantities. Pheophytin a was virtually lacking in allthe samples. Removal rates of pigments, measured by sedimentationtraps, indicated that the degradation of chlorophyll a via chlorophyllidea is a dynamic process that continues during the sedimentationof the phytoplankton particles.     

The dynamics of vertical chlorophyll distribution in an oligomesotropbic lake

The vertical distribution of chlorophyll was examined in relationto physical and chemical parameters in oligomesotrophic LakeSamish, Washington state, during the latter half of a growingseason. A majority of chlorophyll resided in the metalimnionin late July, with metalimnetic populations dominated by diatoms(Cyclotella bodanica) and chrysophytes (Mallomonas caudata,Dinobryon sertularia). A silica gradient appeared to be mostimportant in determining the vertical position of these populations.Later in the season, segregation of nutrients and light, inthe face of a deepening epilimnion, led to transient accumulationsof phytoplankton at the top of the metalimnion. It appearedthat losses due to heterotrophic activities accentuated thelower boundary of this chlorophyll peak.     

Pigment gut contents of copepods and deep phytoplankton maximum in the Western Mediterranean

The feeding activity of Centropages typicus in the Western Mediterraneanwas evaluated during the summer stratification period, whena deep phytoplankton and chlorophyll maximum (DCM) develops.The lack of significant day-night differences in gut contentssuggests continuous phytoplankton ingestion, despite nictemeralmigration of this species. Copepods captured in the DCM hadgreater phytoplankton gut contents, both in nocturnal and diurnalsamples. The observed pattern of phytoplankton ingestion andvertical migration implies copepod-mediated upward transportof nutrients, which might contribute to the maintenance of primaryproduction in the photic layers.     

The presence of chlorophyll b and the estimation of phaeopigments in marine phytoplankton

A reverse-phase h.p.l.c. technique was used to estimate theconcentration of chlorophyll b in phytoplankton cultures, fecalpellets of Calanus pacificus, and suspended paniculate matterfrom the Central North Pacific, Oregon coastal waters, and DabobBay (a temperate fjord in Puget Sound, WA, USA). The purposewas to assess the distribution of this pigment in the euphoticzone and its effect on the fluorometnc estimation of phaeopigments.Analyses of natural waters confirm high chlorophyll b concentrations(median mass ratio of b:a > 0.3) at the depth of the chlorophylla maximum in tropical waters while values for temperate planktonare relatively low (median mass ratio of chl b:a = 0.05) andpatchy. Zooplankton fecal pellets showed a significant enrichmentin chlorophyll b, suggesting grazing as a mechanism to explainhigh concentrations of this pigment at the bottom of the euphoticzone. It is estimated that the presence of chlorophyll b couldcause an average overestimation of phaeopigment concentrationby the fluorometnc technique of 38% between 0 and 200 m in theCentral North Pacific. This effect is more pronounced at thelayer of chlorophyll b maximum (120–140 m). ¹Present address: Marine Biology Research Division, A-002, ScrippsInstitution of Oceanography, La Jolla, CA 92093, USA     

Factors limiting phytoplankton production in a nearshore upwelling area

Phytoplankton productivity was investigated at two nearshoresites in the southern Benguela region during the upwelling season.Changes in biomass and production are discussed in relationto the physical and chemical Status of the water column. Itis suggested that, although light and nutrients affect the productionand biomass of phytoplankton, low phytoplankton biomass in upwellingwater, coupled with the frequency of upwelling, plays a moreimportant role in the overall productivity of the nearshorecoastal zone of upwelling regions where the source water ispoor in chlorophyll a.     

Estimation of phytoplankton photosynthesis using a fluorescence induction technique

An improved method for estimation of phytoplankton photosynthesiswas developed using in-vivo chlorophyll a fluorescence, andwas tested with cultured phytoplankton. The method is basedon a kinetic analysis of the fluorescence induction with andwithout DCMU. Photosynthetic rates were derived from, (i) measurementof fluorescence induction due to the reduction of Q, the primaryelectron acceptor for the photoreaction of photosystem II, and(ii) estimation of Q present in the water sample and of therate of Q reduction. ^*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.     

Phytoplankton and zooplankton development in a lowland, temperate river

The longitudinal and seasonal patterns of plankton developmentwere examined over 2 years in a lowland, temperate river: theRideau River (Ontario, Canada). Following an initial decreasein phytoplankton and zooplankton biomass as water flowed fromthe headwaters into the Rideau River proper, there was an increasein chlorophyll a (chl a) and zooplankton biomass with downstreamtravel. At approximately river km 60, both phytoplankton andzooplankton reached their maximum biomass of 27 µg l^–1(chl a) and 470 µg l^–1 (dry mass), respectively.Downstream of river km 60, the biomass of both planktonic communitiesdeclined significantly despite increasing nutrient concentrationsand favorable light conditions. These downstream declines maybe due to the feeding activity of the exotic zebra mussel (Dreissenapolymorpha) which was at high density in downstream reaches(>1000 individuals m^–2). There was no evidence forlongitudinal phasing of phytoplankton and zooplankton, as increasesand decreases in chl a and zooplankton biomass appeared to coincide.Overall, chl a was best predicted by total phosphorus (R²=0.43),whereas zooplankton biomass was best predicted by chl a (R²=0.20).There was no evidence for significant grazing effects of zooplanktonon phytoplankton biomass.     

Nitrate reductase activity in phytoplankton from the Oslo-fjord, Norway

The activity of the enzyme nitrate reductase (NRA) was measuredin natural populations of phytoplankton in the Oslofjord, Norway,and in culture experiments with 10 different species of phytoplankton.The NRA values from the culture experiments and from the naturalsamples were of the same order of magnitude. The NRA values,when expressed on a chlorophyll a basis, were highly species-variable.In general they were correlated to the nitrogen nutrient concentrationsin the water. In order to use NRA as a guide to an understandingof the assimilation of nitrate, it is important to know thespecies composition of the phytoplankton.     

CHLOROPHYLL AMOUNT AS AN INDICATOR OF MATTER PRODUCTIVITY IN BIO-COMMUNITIES

The data of chlorophyll amounts in diverse bio-communities arecompiled and discussed with reference to matter production. The chlorophyll amount in euphotic zone of lakes and oceanswas less than 1 g/m², mostly less than 0.1 g/m². In phytoplanktonblooms it was ca. 0.1–1 g/m². Large values of 5–20g/m² were obtained in the outdoor mass cultures of Chlorella,in which the high population density and chlorophyll contentof the alga were observed. In terrestrial higher plant communitiesthe chlorophyll amount (ca. 1–10 g/m²) was usually higherthan in aquatic phytoplankton communities. The largest (13.3g/m²) was obtained with an evergreen gallery forest in Thailand.The chlorophyll amount of desert bio-community could be expectedto be as high as that in water blooms. The maximum chlorophyll amount in bio-communities seems to attainup to 20 g/m² when the conditions are favorable. ¹ Dedicated to Prof. H. TAMIYA on the occasion of his 60th birthday. (Received December 25, 1962; )     

Can phaeopigments be used as markers for Daphnia grazing in Lake Constance?

The formation of chlorophyll a degradation products was measuredwith natural phytoplankton from Lake Constance and Daphnia magnaand native Daphnia as grazers in grazing experiments duringspring bloom conditions using high-pressure liquid chromatography(HPLC). Chlorophyll a start concentrations were between 1.2and 16.3 µg l^–1; phaeopigment weights constituted5% of chlorophyll a weight. Only phaeophorbide a was a markerfor Daphnia grazing; concentrations of other phaeopigments (phaeophytina, chlorophyllide a and two unidentified phaeopigments) didnot increase during Daphnia grazing. Conversion efficiencies(chlorophyll a to phaeophorbide a) were between 0 and 43% ona weight basis, and between 0 and 65% on a molar basis. Conversionefficiencies were highest at high grazer density (40 Daphnial^–1) and after a 24 h exposure time. Grazing by microzooplanktonprobably led to the formation of the two unidentified phaeopigments.In Lake Constance, Daphnia density was significantly positivelycorrelated with the phaeophorbide a/chlorophyll a ratio whenit was <5000 Daphnia m^–3. However, when higher Daphniadensities were included in calculations, then Daphnia densitywas positively, but insignificantly, correlated with the phaeophorbidea/chlorophyll a ratio. This suggests that when the level offood per Daphnia is low, then grazing is more efficient withless production of phaeophorbide a and a higher production ofcolourless products.     

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.     

The vertical distributions of chlorophyll and phytoplankton species in the North Pacific central environment

The phytoplankton species in the North Pacific central environmentare known to be distributed into two vertically distinct assemblagesduring most of the year. Key species are defined for each assemblage.The vertical distributions of these key species indicate thatthe increase in abundance of deep species closely parallelsthe increase in chlorophyll a at the top of the chlorophyllmaximum layer. The chlorophyll maximum is comprised of speciescharacteristic of the deep assemblage, with only insignificantnumbers of shallow species.