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.     

Assessment of grazing by the freshwater copepod Diaptomus minutus using carotenoid pigments: a caution

In order to assess feeding selectivity in freshwater zooplankton, we conducted feeding trials using Diaptomus minutus isolated from two Wisconsin lakes. Copepods were fed an algal assemblage comprised of an equal biomass of a centric diatom, a cryptomonad and a coccal green alga. The total amounts of photopigments were tracked using high-performance liquid chromatography. The removals of carotenoids and a-type phorbins (chlorophyll a and phaeopigments) from feeding suspensions were compared with their presence in the guts of animals, in fecal pellets, and in the final suspension. Diaptomus minutus generally removed either the diatoms primarily or all three algal cells equally. These removals were not reflected in gut extracts of the animals, however, where alloxanthin (marker of cryptomonads) was always present, but where fucoxanthin and diadinoxanthin (markers of diatoms) were never observed. Pigment disappearance was variable for total a-type phorbins, but frequently >90% for carotenoids, particularly for fucoxanthin and diadinoxanthin. Phaeophytin a was the major a-type phorbin detected after gut passage. Our results indicate that evaluations of zooplankton grazing which assume that algal carotenoids remain detectable throughout gut passage must be made with substantial caution and that differences in pigment processing are likely to occur among zooplankton species. Furthermore, grazing experiments designed to evaluate decreases of specific pigments in feeding suspension, rather than their appearance in animals' guts, may prove a more valuable approach to understanding the feeding selectivity of copepods.      

Digestion of chlorophylls and carotenoids by the marine protozoan Oxyrrhis marina studied by h.p.l.c. analysis of algal pigments

Algal pigments were measured, by reverse-phase h.p.l.c., duringgrazing experiments with the protozoan Oxyrrhis marina Dujardinon Rhodomonas sp. Chlorophylls and carotenoids were degradedto colourless residues; the rate of degradation of pigmentswas highest in the light. Very little chlorophyll a (5%) wasdegraded to phaeophytin and none to phaeophorbide during grazing.This suggests that phaeopigment concentrations cannot be usedas a measure of algal mortality due to grazing when heterotrophicProtozoa are a component of the grazer community.     

Temporal variability in copepod fecundity during two different spring bloom periods in coastal waters off Plymouth (SW England)

The temporal variation in egg production of the planktonic copepodsCalanus helgolandicus, Temora longicornis and Pseudocalanuselongatus was studied during two different spring bloom periodsin 1989 and 1990 by weekly sampling at two permanent stationsin coastal waters off Plymouth (SW England). Copepod egg productionwas estimated in situ by incubating individual adult femalesin filtered seawater for 24 h (72 h until hatching for P.elongatus)at the field surface temperature. The relationship between copepodegg production rates and: (i) chlorophyll a concentration (totaland >10 µm size fraction), (ii) temperature and (iii)initial copepod gut pigment content was investigated. The springbloom periods were very different in both years, with the occurrenceof a Phaeocystis sp. bloom in 1990, which negatively affectedthe feeding and fecundity of copepods. Egg production ratesin spring 1989 were significantly correlated with chlorophylla concentration (particularly with the >10 µm fraction),field temperature and copepod gut pigment contents. In spring1990, egg production rates were also correlated with copepodgut pigment contents, but no significant correlations were obtainedwith temperature or with chlorophyll a concentration, as a consequenceof the lower egg production rates obtained during the Phaeocystissp. dominance period. These results show that food availabilityis the factor which mainly affects the fecundity of neriticcopepods in short time periods.     

Diverse Responses of Planktonic Crustaceans to Fish Predation by Shifts in Depth Selection and Size at Maturity

We investigated the behaviour and life histories of large zooplankton in the Esch-sur-Sûre reservoir (Luxembourg). We found that the decrease in size at maturity, as well as diurnal refuge in deep waters, were the adaptive responses (concurrently or alternatively) adopted by large zooplankters to cope with the increasing predation risk throughout the summer. Daphnia galeata initiated a more severe trade-off to cope with high summer predation relative to other cladoceran species. Diaphanosoma brachyurum and Daphnia cucullata seem less susceptible to fish predation as indicated by the low alteration of their size structure and their vertical distribution. The copepod Eudiaptomus gracilis took refuge in deep waters and achieved important modifications of its mature size.     

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.     

Phytoplankton Assemblages and Their Dominant Pigments in the Nervion River Estuary

In the Nervion River estuary surface samples were taken from March to September 2003 at six sites covering most of the salinity range with the aim to know the biomass and taxonomic composition of phytoplankton assemblages in the different segments. Nine groups of algae including cyanobacteria, diatoms, dinoflagellates, chlorophytes, prasinophytes, euglenophytes, chrysophytes, haptophytes, raphidophytes and cryptophytes were identified by means of a combination of pigment analysis by high-performance liquid chromatography (HPLC) and microscopic observations of live and preserved cells. Diatoms, chlorophytes and cryptophytes were the most abundant algae in terms of cells number, whereas fucoxanthin, peridinin, chlorophyll b (Chl b) and alloxanthin were the most abundant auxiliary pigments. Based on multiple regression analysis, in the outer estuary (stations 0, 1, 2 and 3) about 93% of the chlorophyll a (Chl a) could be explained by algae containing fucoxanthin and by algae containing Chl b, whereas in the rest of the estuary most of the Chl a (about 98%) was accounted for by fucoxanthin, Chl b and alloxanthin containing algae. The study period coincided with that of most active phytoplankton growth in the estuary and fucoxanthin was by far the dominant among those signature pigments. Several diatoms, chrysophytes, haptophytes and raphydophytes were responsible for fucoxanthin among identified species. Besides, dinoflagellates with a pigment pattern corresponding to chrysophytes and type 4 haptophytes were identified among fucoxanthin-bearing algae. Cryptophytes were the most abundant species among those containing alloxanthin. The maximum of Chl b registered at the seaward end in April coincided with a bloom of the prasinophytes Cymbomonas tetramitiformis, whereas the Chl b maxima in late spring and summer were accounted for by prasinophytes in the middle and outer estuary and by several species of chlorophytes in the middle and inner estuary. Other Chl b containing algae were euglenophytes and the dinoflagellate Peridinium chlorophorum. Dinoflagellates constituted generally a minor component of the phytoplankton.     

Pigment destruction by Calanus pacificus: impact on the estimation of water column fluxes

Measurements of gut pigment made prior to fecal pellet productionin Calanus pacificus females and CV copepodites suggest that(i) chlorophyll a and/or its pheopigment derivatives are degradedinto molecules that are not detectable by the standard fluorometrictechnique; and (ii) the percentage of ingested chlorophyll awhich degrades into fluorometrically undetected molecules isnot constant. Thus, measurements of chlorophyll and pheopigmenta in the guts of zooplankton can only yield minimum estimatesof in situ grazing rates. Estimates of the vertical flux ofprimary particulates based on chlorophyll and pheopigment abudgets may also be underestimated.     

Diel activity of Gammarus pulex (Crustacea) in a South Swedish stream: comparison of drift catches vs baited traps

The benthos of a perennially ice-covered Antarctic lake, Lake Hoare, contained three distinct signatures of lipophilic pigments. Cyanobacterial mats found in the moat at the periphery of the lake were dominated by the carotenoid myxoxanthophyll; carotenoids: chlorophyll a ratios in this high light environment ranged from 3 to 6.8. Chlorophyll c and fucoxanthin, pigments typical of golden-brown algae, were found at 10 to 20 m depths where the benthos is aerobic. Anaerobic benthic sediments at 20 to 30 m depths were characterized by a third pigment signature dominated by a carotenoid, tentatively identified as alloxanthin from planktonic cryptomonads, and by phaeophytin b from senescent green algae. Pigments were not found associated with alternating organic and sediment layers. As microzooplankton grazers are absent from this closed system and transformation rates are reduced at low temperatures, the benthos beneath the lake ice appears to contain a record of past phytoplankton blooms undergoing decay.     

Gut evacuation rates and ingestion rates of Eudiaptomus graciloides measured by means of the gut fluorescence method

Journal of Plankton Research, 8, 973–983, 1986 FIg. 2. Time-dependent changes in the gut content (percentageof initial ng pigment) of E. gro.ciloides at different temperaturesunder simultaneous feeding. Fig. 4. The relationship between instantaneous evacuation rateand temperature of E. graciloides. The regresston equation forfeeding animals: y = 0.0044 e(0.141 ) (r² = 0.90). For comparisonthe results of non-feeding animals are indicated with open circles.     

Enzymatic degradation of chlorophyll a by marine phytoplankton in vitro

The enzymatic degradation of chlorophyll a and the formation of chlorophyllide a, phaeophytin a, and phaeophorbide a were detected in vitro in several species of marine phytoplankton. Loss of phytol and Mg²⁺ were found to be catalysed by chlorophyllase and a magnesium-releasing enzyme, respectively. The activities of the two enzymes could be distinguished from each other by inhibiting with Mg²⁺ and/or p-chloromercurobenzoate. Both enzymes are activated by cell disintegration. Degradation products were not detected spectrophotometrically in vivo. Additionally, in some species, chlorophyll a was degraded to products which do not absorb visible light.     

Degradation of diatom chlorophyll to colourless,non-fluorescing compounds during copepoo grazing

During grazing of the copepodTemora longicornis onThalassiosira eccentrica phaeophorbide-a and several phaeophytins-a appeared in the culture vessels. However, a variable portion of the diatom's chlorophyll-a was degraded to substances that could not be detected either spectrophotometrically or fluorometrically. Chlorophyll-c and fucoxanthin were also lost during diatom cell transit throughTemora's gut. The variability between experiments in the destruction of chl.a to colourless residues is ascribed to differences in coprophagy and in transit time through the copepod gut, processes related to filtering rate and the ambient food concentration. Because the pigment loss is variable, grazing pressure cannot be estimated by simply recording bulk phaeopigment concentrations. The net result of the appearance of a fucoxanthinol-like pigment while all other pigments disappeared is a gradual carotenoid enrichment of the samples during grazing.     

Study of selective feeding in the marine cladoceran Penilia avirostris by HPLC pigment analysis

Food selection by the marine cladoceran Penilia avirostris was studied in the field by HPLC analysis of phytoplankton marker pigments and in the laboratory by microscopic measurement of cell removal. Comparison between pigment composition in natural phytoplankton and in P. avirostris showed that P. avirostris preferred diatoms, cryptophytes and chlorophytes, and ignored prymnesiophytes and dinoflagellates. Peridinin, the marker pigment for dinoflagellates was found in P. avirostris only when the dinoflagellate populations were dominated by Prorocentrum. Pigment degradation rates ranged from 13.73% for alloxanthin to 36.62% for chlorophyll a. Clearance rates measured in the laboratory provided further evidence of strong preference for diatoms and cryptophytes, and avoidance of dinoflagellates. Microscopic counts suggested that P. avirostris was feeding on prymnesiophytes, although ingestion of prymnesiophytes could not be confirmed by pigment analysis.     

Phytoplankton pigments at the Weddell-Scotia confluence during the 1993 austral spring

During a 1993 austral spring cruise, a complex biomass was encounterednear South Orkney Island that ranged from a low-biomass, Chaetocerostortissimus assemblage south of the front towards the ice edge,to a high-biomass, Thalassiosira gravida-dominated assemblageat the northern edge. The maximum levels of chlorophyll (Chl)a (up to 6 mg m^–3) were higher than those observed inprevious high-performance liquid chromatography-based studiesof pigments in the pelagic Southern Ocean. The non-photosyntheticpigment chlorophyllide a comprised up to 75% of the chlorophyllouspigments in the southern assemblage, but < 5% in the northernassemblage. Concentrations of the xanthophylls diadinoxanthin(DD) and diatoxanthin (DT), used as indicators of mean irradiance,indicated low-light-adapted populations. Low-light DD + DT/Chla ratios in surface waters indicated that vertical mixing limitedphytoplankton residence time in the near-surface layer, andthus limited exposure to maximum irradiance. Deck incubationsof natural assemblages indicated that the dark epoxidation reaction(i.e. the return of DT to DD) was a two-step reaction with theinitial rate being more rapid (t^1/2 = 9.5 min) than the second(t^1/2 = 55 min). Fucoxanthin, a major diatom pigment, was morestable chemically in the water column than Chl a, and the verticalprofiles of fucoxanthin followed those of chlorophyllide a insome cases. The formation and apparent stability of chlorophyllidea and fucoxanthin are important considerations when estimatingphotosynthetically active biomass over large regions of theocean.     

Copepod feeding study in the upper layer of the tropical South China Sea

The South China Sea (SCS) is the world’s largest marginal sea being notable for vertical mixing at various scales resulting in a sequence of chemical and biological dynamics in surface waters. We investigated the ingestion, gut content, evacuation and clearance rates of copepods collected from six stations (including a South East Asia Time Series station) along a transect line in the tropical of a SCS cruise during September 27, 1999 to October 2, 1999. The goal of the present study was to understand the feeding ecology of copepods in the upper water layers (0–5 m) of the northern SCS during autumn. We measured the gut pigment contents of 33 copepod species by the gut fluorescence method. The gut chlorophyll a values of most small size copepods (<1 mm) were lower than 1.00 ng Chl a individual⁻¹. The highest gut pigment content was recorded in Scolecithrix danae (7.07 ng Chl a individual⁻¹). The gut pigment contents of 33 copepod species (including 70 samples and 1,290 individuals) estimated is negatively correlated with seawater temperature (Pearson correlation r = −0.292, P = 0.014) and is positively correlated with the chlorophyll a concentration of ambient waters (Pearson correlation r = 0.243, P = 0.043). Mean gut pigment content, ingestion and clearance rates (from 80 samples and 1,468 individuals) show that larger copepods (>2 mm) had significantly higher values than medium sized copepods (1–2 mm) and smaller sized copepods. The present study shows that the performance of feeding on phytoplankton was variable in different sized copepod groups, suggesting that copepods obtained in the tropical area of the southeastern Taiwan Strait might be opportunistic feeders.     

Influence of phytoplankton composition and stratification degree on gut pigment content of Ceriodaphnia sp. at dawn

Very short-term feeding activity of the cladoceran Ceriodaphniasp. was investigated in situ in a eutrophic reservoir in thesouth of Spain, using fluorimetric analysis of the gut pigmentcontent in periods when the water column was relatively mixedor strongly stratified. The results obtained in the mixed watercolumn showed a clear increase in gut pigment content at dawn,a period sampled with high frequency. The accumulation of thecladoceran at the depth of maximum concentration of phytoplankton,and the high gut pigment concentration in cladocerans at thatdepth just after dawn, suggested active feeding of Ceriodaphniaon phytoplankton at that time. During stratification, the abundanceof Ceriodaphnia was higher, but the gut pigment contents werevery low and they did not reflect any clear feeding patterns,with either time or depth. Changes in phytoplankton concentrationand composition between the relatively mixed and the stratifiedwater column suggest a shift in feeding activity from herbivorousto.     

Gut pigment destruction by the copepod Acartia clausi

The extent of chlorophyll (Chi) a degradation into colorlessproducts by the copepod Acartia clausi was determined by measuringgut fluorescence prior to fecal pellet production. The efficiencyof pigment degradation of animals fed at different concentrationsof Thalassiosira weiss-fiogii varied between 30.56 and 94.05%,and the quantity of pigments lost during gut passage was directlyrelated to total ingestion. The role of feeding history in pigmentdestruction was tested. The fraction of ingested Chi a degradedby copepods acclimated to a high food concentration (15 ng Chla ml^–1) was greater than the fraction degraded by thoseacclimated to a low food concentration (1.5 ng Chl a ml^–1).The percentage of ingested Chl a that A.clausi transformed intofluorometrically undetected compounds was not constant. At present,knowledge of the pigment destruction process indicates thatChi a and Chi --derived pigments are not considered useful quantitativetracers of the feeding activity of copepods. The results ofthis study suggest a reassessment of the application of thegut fluorescence method to evaluate grazing activity of thecopepod A.clausi     

Pigment Composition of a Novel Oxygenic Photosynthetic Prokaryote Containing Chlorophyll d as the Major Chlorophyll

The principal pigment found in the majority of oxygenic photosyntheticorganisms is known to be chlorophyll a. However, we isolateda new oxygenic photosynthetic prokaryote that contained chlorophylld as a predominant pigment with chlorophyll a being a minorpigment. Chlorophyll d had previously been noted but its naturaloccurrence and function remained unclear. Cells of the new prokaryotehad an absorption maximum at red region of 714–718 nmdue to chlorophyll d absorption, but no characteristic absorptionpeak of chlorophyll a around 680 nm was observed. Chlorophylld of the new organism was identified spectrophotometricallyin several solvents and its chemical structure was confirmedby NMR and FABMS analysis. The cell also contained a chlorophyllc-like pigment, zeaxanthin and a-carotene but not chlorophyllb and ß-carotene. The content of chlorophyll d accountedfor more than 2% of the cell dry weight, while the content ofchlorophyll a was less than 0.1%. The chlorophyll a/d ratioremained between 0.03 and 0.09 under different culture conditions.The light absorption characteristics and the high content ofchlorophyll d along with the small content of chlorophyll aindicated the existence of a new light utilization mechanisminvolving chlorophyll d. (Received October 7, 1996; Accepted December 16, 1996)     

Diel periodicity of Subantarctic copepods: relationships between vertical migration, gut fullness and gut evacuation rate

A zooplankton community in the Polar Frontal Zone north of SouthGeorgia was sampled for 5 days in February 1994. Feeding ofvarious copepodite stages of six copepod species was assessedby a series of gut fluorescence/gut evacuation experiments.Feeding periodicity was compared to vertical distribution andmigration patterns, as revealed by Longhurst Hardy PlanktonRecorder (LHPR) and ring net catches. Despite chlorophyll alevels reaching only 0.8 mg m^–3 and daily carbon rationsbased on phytoplankton intake being low, feeding of all thecopepods was restricted mainly to the 8 h night-time perid.During the daytime, the epipelagic community was verticallydispersed within the top 100 m. At night, upward migration bymost species led to a convergence of almost all zooplanktersin the upper half of the surface mixed layer. However, large-amplitudevertical migrations which crossed the thermocline were performedonly by Metridia lucens and Pleuromamma robusta. Although feedingby both migrants and non-migrants was mainly at night, therewas no diel signal in gut evacuation rate. The gut evacuationrates of the nine copepod species and stages differed significantly(5-fold) and were negatively related to the extent of theirdiel vertical migration. The long gut-passage times of the migratoryspecies, M.lucens and P.robusta, would have allowed them timeto defaecate some of the food eaten near the surface at depth,contributing to an active carbon transfer out of the mixed layer.However, their scarcity at this site meant that their grazingcomprised only -<1.4% of total copepod ingestion, so theircombined gut flux is likely to have been negligible (-0.4 mgcarbon day^–1).     

Variations in biochemical parameters of Heterocapsa sp. and Olisthodiscus luteus grown in 12:12 light:dark cycles II. Changes in pigment composition

Photosynthetic pigment composition was studied in batch cultures of Heterocapsa sp. and Olisthodiscus luteus growing exponentially in a 12:12 light:dark cycle. Both species divided in the dark. The synthesis of pigments was continuous for both species. However for chlorophyll c and peridinin, in Heterocapsa sp., and chlorophyll c and fucoxanthin, in O. luteus, (pigments belonging to light harvesting complexes) the synthesis was significantly higher during the light period. Concentrations per total cell volume (TCV) of chlorophyll a, chlorophyll c, peridinin and diadinoxanthin in Heterocapsa sp., and chlorophyll a, chlorophyll c, fucoxanthin and violaxanthin in O. luteus, showed a maximum at the onset of light and decreased during the light period. The values of the chlorophyll a:chlorophyll c, chlorophyll a:peridinin and chlorophyll a:fucoxanthin ratios are compared with data reported in the literature.