The production and ingestion of faecal pellets by nauplii of marine calanoid copepods

The downward transport of organic matter as zooplankton faecalmaterial is influenced by copepods which fragment, ingest andrecycle some of the pellet contents. Most of this activity hasbeen attributed to the later copepodite stages and the adults,but little is known about the role of nauplii. Stage-relateddefaecation rates during the naupliar development of two speciesof copepod, Calanus helgolandicus and Pseudocalanus elongatus,were quantified in a series of laboratory experiments. The productionof faecal material commenced soon after the appearance of theNIII in both species and increased throughout naupliar development.The causes of the increase were the formation of larger pelletsby later stages in Calanus and an increased rate of productionby Pseudocalanus. Calanus nauplii, when supplied with algalfood at concentrations that would support full naupliar development,ingested or broke up the pellets of the smaller Pseudocalanusspecies at rates of 1.15 pellets nauplius^–1 h^–1This consumption increased to 2.96 pellets nauplius^–1h^–1 when the concentration of algal food was reduced toa limiting level. Pseudocalanus was not able to consume thepellets of Calanus. Ingestion of Pseudocalanus faecal pelletsby Calanus could supply a nutritional benefit to a food-limitednauplius.     

Developmental rates of the copepods Calanus australisand Calanoides carinatus in the laboratory, with discussion of methods used for calculation of development time

The copepods Calanus australis and Calanoides carinatus developedfrom egg to adult in 20.3 and 18.3 days respectively at 15.5?Cand 16.0 and 12.0 days at 19.5?C. For both species the durationof the first two naupliar stages was short, <1 day. The thirdnaupliar stage was prolonged in C.australis but short in C.carinatus. Isochronal development was approximated from fourthnauplius through third copepodite; fourth and fifth copepoditestages were relatively long. Only female C.australis were produced;60% of the C.carinatus were female. Equiproportional developmentwas found for C.carinatus but not for C.australis. For bothspecies the variance in stage duration was established at thethird nauplius then remained unchanged through the older stages.In calculating developmental rates several commonly used methodswere compared. The method of ‘first appearance of an individualof a given stage’ and ‘mean time of appearance ofa stage’ both yielded estimates of development time thatwere 10–15% faster than the more commonly used methodof ‘median development time’. We suggest that allfuture work on developmental rates of copepods should use astandardized method for the calculation of stage-specific developmentalrates and that ‘median development time’ be themethod of choice. ³Present address: Marine Sciences Research Center, SUNY, StonyBrook, NY 11794-5000, USA     

The escape behavior of marine copepods in response to a quantifiable fluid mechanical disturbance

The threshold shear values needed to elicit the escape reactionto a quantifiable fluid mechanical disturbance were comparedbetween five free-swimming oceanic copepod species. The resultsindicate a significant difference in the threshold for differentspecies of copepods and between different age groups withina single species. In general, animals captured from more energeticregimes required a higher threshold than those captured frommore pacific locations. Labidocera madurae required the highestshear values with 51.5 s^–1 for 50% of the animals testedto elicit an escape reaction (S₅₀). Acartia tonsa and Euchaetarimana, in contrast, were behaviorally the most sensitive requiringan S₅₀ of only 1.5 and 4.1 s^–1, respectively, to initiatean escape reaction. Pleuromamma xiphias and Oithona requiredintermediate shear values with an S₅₀ of 7.2 and 8.1 s^–1.When compared to literature values, the threshold needed toelicit an escape reaction was consistently higher than averageenvironmental shear values. Threshold shear values also variedsignificantly with developmental stage. Naupliar stages of A.tonsarequired greater than six times the S₅₀ value required by adultsof the same species. This suggests that the higher vulnerabilityto predation of naupliar stages of copepods may not only reflectinferior escape strength, but may also result from the higherthreshold needed to elicit an escape reaction. This study supportsthe hypothesis that selective feeding patterns exhibited bypredators of copepods may be the result of the differentialbehavioral sensitivities of different species and developmentalstages of copepods.     

The effect of salinity and temperature on the larval development of Mithrax caribbaeus Rathbun, 1920 (Brachyura: Majidae) reared in the laboratory

Larvae of Mithrax caribbaeus were reared in the laboratoryin a factorial experiment employing three temperatures (22,25 and 28°C) and three salinities (32, 35 and 38). Survivaland duration of larval stages were recorded. Ovigerous femalesof M.caribbaeus were collected from the south-eastern coastof Margarita Island, Venezuela, and maintained in individualaquaria until hatching. Eggs from three of the females hatchedin the laboratory. Larvae from each hatching were subdividedinto groups of 10 and reared in plastic bowls containing 200ml filtered and UV-irradiated sea water at different temperature–salinitycombinations. Larvae were transferred daily to clean bowls withnewly hatched Artemia nauplii, and the number of molts and mortalitywithin each bowl was recorded. Complete larval development ofM.caribbaeus occurred under all experimental conditions. Salinityhad the greatest effect on percentage survival of each larvalstage and complete development up to the first crab stage. Thefirst zoeal stage exhibited the highest survival rate. Maximumsurvival for this stage occurred at 25°C, 32–35. Survivalin the second zoeal stage and the megalopa was affected onlyby salinity. Effects of temperature and salinity on survivaldecreased with advance in development. The duration of the twozoeal stages, the megalopa, and development to the first crabstage showed a gradual reduction with increasing temperature.Salinity showed an effect on the duration of zoeal stages butnot on the megalopal stage. Development from hatching to thefirst crab stage required 8–18 days, depending on thetemperature–salinity combination, and was inversely relatedto temperature, averaging 14.3 days at 22°C, 11.8 days at25°C and 9.2 days at 28°C.     

Herbivorous nutrition of Cyclops vicinus: the effect of a pure algal diet on feeding, development, reproduction and life cycle

Nutritional requirements, functional response, development andreproduction of Cyclops vicinus were studied with exclusivelyalgal food. Phytoflagellates were found to be adequate foodresources for both juvenile development and egg production.Ingestion measurements were performed with Chlamydomonas reinhardii.The functional response data give evidence for low feeding efficiency,especially for the naupliar stages. A difference between naupliiand older instars was also found in their quantitative foodneeds. A higher threshold food concentration was observed fornaupliar development (0.4 mg C l^–1) than for copepoditedevelopment (0.2 mg C l^–1). The calculation of assimilationefficiencies suggests that the high food requirements are dueto low specific ingestion rates rather than poor assimilationefficiency. Development time decreased as algal density increasedand males developed more quickly than females at all food concentrations.Body size and carbon content increased with increasing foodconcentration. Continuous egg production was observed abovea food concentration of 0.5 mg C l^–1. The results haveimplications for the life cycle of C. vicinus. Summer diapauseis interpreted as a strategy to avoid starvation of the juvenilestages. The facultative herbivory of the adults might be anadvantage when competing against other more carnivorous cyclopoidcopepods. ¹Present address: Agricultural University of Wageningen, Departmentof Mathematics, Dreijenlaan 4, NL-6703 HA Wageningen, The Netherlands     

Cannibalistic feeding behavior of the brackish-water copepod Sinocalanus tenellus

Cannibalistic feeding behavior of the brackish-water copepodSinocalanus tenellus was examined in the laboratory using CI-II,CIII-IV and CVI female as predators and NI-II, NIII-IV, NV-VIand CI-II as prey. In each prey-predator combination, the ingestionrate increased with increasing prey density to an asymptoticvalue. Cannibalism took place even when phytoplankton was availableas an alternative food supply. Based on a daily ration, theoptimal prey stages for CVI females, CIII-IV and CI-II are NI-VI,NI-IV and NI-II respectively. Under average, natural prey density(10 nauplii l^–1), S tenellus can achieve only a smallfraction (max 9%) of the daily minimum food requirement by cannibalisticfeeding. However, the impact of cannibalism on naupliar survivorshipcan be significant. When adult females occur at a density of10 l^–1, the mortality due to cannibalism attains 99.2%during the naupliar stages.     

The influence of food resources on the development, survival and reproduction of the two cyclopoid copepods: Cyclops vicinus and Mesocyclops leuckarti

Laboratory experiments were conducted to determine whether thetwo cyclopoid copepods. Cyclops vicious and Mesocyclops leuckaru.exploit the same food resources. The food requirements of juvenilesof the two cyclopoid copepods were investigated. Moreover, theimportance of algae for the predaceous adults was studied. Naupliiof both M leuckaru and C.vicinus successfully developed intocopepodites when fed the motile algae Chlamydomonas reinhardtii.Chlamydomonas sphaeroides and Cryptomonas sp. Threshold foodconcentrations for naupliar development varied between offeredalgae and between the two cyclopoid species. The food thresholdfor successful naupliar development, when reared on C.reinhardui,was lower for M.leuckarti (0.3 mg C 1^–1) than for C.vicinus(0.5 mg C l^–1) whereas a similar food threshold was foundusing Cryptornonas sp (0.3 mg C ^–1) and C.sphaeroides(<0.2 mg C 1^–1), Naupliar development time was inverselyrelated to food concentration. Food required for copepoditedevelopment differed for the two cyclopoid species. Cyclopsvicinus was able to develop to the adult stage on a pure dietof any one of the three algal species. whereas M.leuckarti requireda prey supply of the rotifer Brachionus rubens. Food composition.i.e. algal species, algal concentration and rotifer abundance,influenced copepodite survivorship of both cyclopoids and wasalways higher in the presence of B.rubens. Under similar foodconditions, mortality was higher for M.leuckarti than for Cvicinus. Mesocyclops leuckaru females were very dependent onanimal food. The predation rate of M.leuckaru was not lowerin the presence of algae. Egg production of M.leuckarti waslow on a pure algal diet and significantly higher when B rubenswas present. The results were used to discuss the life cyclestrategy and the possibility of exploitative competition ofthe two cyclopoid copepods.     

Effect of photoinhibition on algal photosynthesis: a dynamic model

Recent evidence from algal physiology and molecular biologyconfirms that photoinhibition is directly related to D1 proteindamage and recovery, and D1 protein damage leads to a decreasein electron transfer or an increase in turnover time of theelectron transfer chain. In this study, the turnover time ofthe electron transfer chain is defined as a function of therelative concentration of D1 protein in reaction centre II andthe photoinhibition processes due to D1 protein degradationare incorporated into a model of photosynthesis, initiated byDubinsky et al. (Plant Cell Physiol., 27, 1335–1349, 1986)and developed by Sakshaug et al. (Limnol. Oceanogr., 34, 198–205,1989). D1 protein damage is assumed to be both light and D1protein concentration dependent, and to be proportional to thecross-section of PSII (     

Structural studies on the tri- and tetrasaccharides isolated from porcine intestinal heparin and characterization of heparinase/heparitinases using them as substrates

We prepared a series of oligosaccharides from porcine intestinalheparin after extensive digestion with a mixture of Flavobacteriumheparinase as well as heparitinases I and II. Previously, wereported the structures of the two glycoserines derived fromthe carbohydrate-protein linkage region [Sugahara et al., J.Biol. Chem., 267, 1528–1533 (1992)] and three tetrasaccharidesderived from the antithrombin III-binding site [Yamada et al.,J. Biol. Chem., 268, 4780–4787 (1993)]. In this study,we determined the structures of 10 other tetrasaccharides anda trisaccharide by enzymatic digestion, fast atom bombardmentmass spectrometry and 500-MHz ¹H NMR spectroscopy. These tetrasaccharidesshare the common disulphated structure,     

Development of Eodiaptomus japonicus Burckhardt (Copepoda, Calanoida) reared on different sized fractions of natural plankton

The nutritional value of different sized fractions of naturalplankton was investigated for the growth of Eodiaptomus japonicusBurckhardt by comparing the development of its naupliar andcopepodid stages fed on differentially fractionated planktonicassemblages of a eutrophic pond, at 20°C. Water filteredthrough a 0.8 µm Nuclepore filter, containing mainly smallcoccoid bacteria (0.45–0.6 µm in cell diameter),at a concentration of 82.7 µg C 1^–1 could not supportthe development of E.japonicus. The 3 µm filtered water,containing bacteria and picoalgae. at a total concentrationof 259 µg C 1^–1, supported development but not eggproduction. The 20 µm filtered water, containing bacteria,picoalgae and large algae, at a total concentration of 2600µg C 1^–1, supported rapid development of the juvenilesand continuous egg production by the adults. The separated 3–20µm fraction, containing only large algae, could not supportthe development at concentrations of 131 and 196 µg C1^–1. However, the same rapid development of the juvenilesand continuous egg production by adults occurred at all of thetested concentrations between 261 and 3920 µg C1^–1of the large algae. The results suggest that E.japonicus favoursalgae larger than 3 µm during its complete lifespan, andthat the threshold food concentration for its development variesbetween 200 and 250 µg C 1^–1.     

Post-embryonic developmental rates as a function of food type in the cyclopoid copepod, Mesocyclops thermocyclopoides Harada

We studied under laboratory conditions the post-embryonic developmentalrates (from nauplii to adult stage) of Mesocyclops thermocyclopoides,commonly found in the eutrophic and hypertrophic local ponds,in relation to different food types. Nauplii hatched from theeggs collected from laboratory-maintained mass cultures werereared at 25l.5C, using the following test diets: bacteria(Klebsiella aerogenes), cyanobacteria (Microcystis aeruginosa,as single cells and in the form of colonies), green algae (Chlorella,Scenedesmus and Chlorogonium), ciliates (Tetrahymena) and rotifers(Brachionus angularis). The developmental rates were fastest,and the proportion of nauplii reaching the copepodid stage andthat of copepodids reaching the adult stage the highest, withChlorogonium, followed by ciliates and rotifers. Developmentwas incomplete with bacteria and Microcys-tis.Ttie algae Chlorellaand Scenedesmus supported complete post-embryonic developmentonly when cultured in media enriched with beef extract, bactotryptoneand yeast extract. With rotifers as the exclusive food, >70%of the nauplii reached the copepodid stage, and >85% of thecopepodids the adult stage. The ratio (D_c/D_n) of copepodid duration(D_c) and naupliar duration (D_n) was significantly differentfrom 1.0, indicating the absence of isochronal development inM.thermocyclopoides. Our results show that post-embryonic developmentin this species is not possible with certain algal diets, probablybecause of nutritional deficiencies, and that the later stagenauplii are capable of capturing rotifers and utilizing themas food to complete their development to the adult stage.     

Temperature-dependent stage durations of Irish Sea Nephrops larvae

The relationships between temperature and stage duration ofthe three zoea stages of Irish Sea Nephrops norvegicus wereinvestigated by culturing individual larvae at five differenttemperatures. The relationships are required for estimatingdaily larval production from survey data on abundance. The survivalof cultured larvae was relatively high. Estimates of the temperature–stageduration relationships from data of individual larvae were differentfrom a previous Irish Sea study. However, using the averageduration at each temperature gave relationships that were notsignificantly different from previous studies in the Irish Seaand Clyde. No larvae were successfully reared to stage IV (post-zoea)at lower temperatures, so it was impossible to assess the relationshipbetween stage III larval duration and temperature.     

Motion behavior of nauplii and early copepodid stages of marine planktonic copepods

The goal of this study was to quantify periods of activity andvelocities of late naupliar and early copepodid stages of planktoniccopepods occurring regularly on the southeastern continentalshelf of the USA. We obtained quantitative information on eightspecies, including adult females of Oithona plumifera. All studieswere conducted at food concentrations near or above satiationlevels. Activities ranged from 0.85% (adult females of O.plumifera)to 100% of time (nauplii and copepodids of various calanoidspecies). Motion velocities (excluding escape motion) coveredmore than one order of magnitude: from 0.39 mm s^–1 fornauplii of Temora stylifera to 5.24 mm s^–1 for naupliiof Oncaea mediterranea. Ranges of activities of species rangefrom occasional for early juveniles to adult females of O.plumiferato 100% for the same range of T.stylifera, the latter creatinga feeding current from N III onwards, the former not at all.Of notable interest is Centropages velificatus which moves intermittentlyas a late nauplius, continuously as an early copepodid and intermittentlyas an adult. All observed calanoid late nauplii and copepodidsmove in three dimensions, excluding copepodids of the shelfbreak/oceanicParacalanus aculeatus. The results indicate not only significantdifferences in motion behavior between cyclopoids and calanoids,but also between calanoid species. Yet, some calanoid speciesshow little ontogenetic changes at all.     

The year cycle of Eudiaptomus vulgaris (Schmeil, 1896) (Copepoda,Calanoida) in a small,acid water body during 1973. Development in the natural habitat and relationships between temperature and duration of development stages

The development of a population of Eudiaptomus vulgaris (Schmeil, 1896) in the Meeuwenven, a shallow acid guanotrofic moorland pool, has been described during one year. The population hibernates as copepodite 5 stages, adults and, to a small extent as naupliar stages N1, N2 and N3 (which could not develop further at low temperatures in autumn). In spring the population development starts at temperatures above 10°C and shows 3 or 4 pulses a year. An attempt has been made to explain seasonal changes in the size of adult males and females and in the sex ratio.In order to establish the duration of the various development stages, the animals have been cultured at different temperatures under illumination with 2000 Lux at a daylength of 14 hours. An adequate quantity of food from the natural habitat was available.Total egg development and total naupliar and copepodite development have been compared with the results of other workers, especially with those from Eckstein (1964), who studied Eudiaptomus vulgaris in the deep Schluchsee. The duration curves do not differ markedly with those of Eckstein and are strongly temperature dependent.The relation between the development times of the various stages with temperature can be generally expressed as parabolic regressions of the type D = a + b₁T + b₂T², the C₅ and adult stages being the only exception at higher temperatures. Comparison of the relative duration of the stages at different temperatures did show that younger stages can take a larger share of the total development time at lower temperatures, stage N6 being the most temperature-sensitive.     

Food size selectivity of Artemia franciscana at three developmental stages

Food size selectivity was examined in Artemia franciscana metanaupliiat three different developmental stages. Clearance rates weredetermined in short-term experiments either by measuring thedecrease in concentration of live particles and plastic beads,or by measuring the radioactivity accumulated in animals thatgrazed ¹⁴C-labelled live particles. The maximum clearance rateofA.franciscana metanauplii increased during development andwas measured at 50–63 µl ind.^–1 h^–1,254 µl ind.^–1 h^–1 and 1.48–2.10 ml ind.^–1h^–1 in 2-, 4- and 7-day-old metanauplii, respectively.A preference for particles with a diameter of 4–8 µmwas observed at all three developmental stages. The abilityof A.franciscana metanauplii to graze bacterial particles wasalso demonstrated, although the efficiency in grazing such smallparticles was low compared to microalgae (28, 20 and 9% of themaximum clearance rate in 2-, 4- and 7-day-old metanauplii,respectively). Electron microscopy showed that the inter-setulardistance in antennae and thoracopods was 0.20 ± 0.07,0.16 ± 0.05 and 0.18 ± 0.04 µm in 2-, 4-and 7-day-old metanauplii, respectively, and accordingly independentof stage.     

Estimate of the effects of ultraviolet radiation on the mortality of Artemia franciscana in naupliar and adult stages

The impact of different doses of artificial ultraviolet (UV) radiation on the growth stages of a marine zooplankton was investigated using laboratory microcosms. Mortality percentages of naupliar and adult samples of Artemia franciscana were recorded in relation to different UV doses (single exposure: 75, 150, 300, 600, 1,200, 2,400, 3,900, 7,800 J m^–2) at specific observation times after exposure (24, 48, 72, 96 and 120 h). The relationship between mortality percentage and UV dose showed significant differences in relation to the zooplankton growth stage. The elevated susceptibility of the naupliar samples to UV radiation is described through a mortality model based on a logistic equation. The data analysis shows that the slope of mortality versus dose remains the same for the two growth stages while the lethal dose in the naupliar stage was 3.3 smaller than that determined for the adult stage. The slope of the UV mortality rate versus post-incubation time was found to be significantly different (P<0.05) at low UV doses for the two life stages examined, i.e. naupliar and adult. The lower value of LD₅₀ in naupliar stages compared to that for adults confirms that in the early growth stage this marine zooplankton is more susceptible to UV radiation.     

The complete postembryonic development of Tropodiaptomus informis Kiefer, 1936 (Copepoda:Calanoida) reared in the laboratory

The complete postembryonic development of Tropodiaptomus informiscomprises six naupliar and six copepodid stages of which thelast is the adult. Of all the characteristics, it is the shapeand number of setae on the terminal segments of the antennulesin the late naupliar stages, and the shape and setae of exo-and endopods of the fifth legs from copepodids III to V whichare most useful in constructing relationships among eight diaptomidsreared.     

Characteristics of Chlorophyll Formation in the Green Alga Golenkinia

Cells of the alga Golenkinia are bleached by growth in darknessin media containing sodium acetate. Re-greening of these cellsis light dependent; neither glucose nor intermediates of chlorophyllsynthesis can substitute. The amount of chlorophyll synthesizedis proportional to the light intensity between darkness and1,000 lux and to the duration of the exposure. Initially, onlychlorophyll a is synthesized. After 9–12 hr illumination,formation of chlorophyll b and carotenoids begins. Chlorophyllproduction apparently occurs in two stages: (1) the first 12–16hr of greening. This stage is sensitive to cyanide, azide oranaerobiosis and relatively resistant to DCMU. (2) the second16–24 hr of greening. This stage is sensitive to DCMUand relatively resistant to inhibitors of respiration. Glucosestimulates greening in both stages. The metabolic requirementsof chlorophyll synthesis are discussed. (Received December 17, 1980; Accepted June 25, 1981)     

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.     

Size, composition and distribution of particles related to wind induced resuspension in a shallow tropical lagoon

The size, composition and distribution of particles in the watercolumn were surveyed in a shallow area (1 m depth) of a tropicallagoon (Cte d'Ivoire) during a sequence of wind-induced resuspension.Water samples were collected hourly near the surface duringone tidal cycle. Three characteristic periods were distinguished:a calm period with low wind speed (average 1.2 m s^–1 awindy period with wind speed >3 m^–1 s (range between4 and 6 m s^–1) inducing sediment resuspension and a relaxationperiod during the decrease of wind velocity. From the analysisof several parameters (particle size and volume, bacteria. pico-and nanophytoplankton, ciliates and detritus), sediment resuspensioncaused a regular injection of particles from the bed. The finestparticles (1.5–6 µm: chlorophytes such as Chiorellaspp., picocyanobacteria such as Synechococcus) were the firstto be affected by wind-induced turbulence, whereas large particles(6–12 µm: diatoms. cyanobacteria such as Lyngbiaspp.) were dispersed into the water column at the highest windspeed. The fate of the different seston components differedaccording to their size. Therefore, wind-induced resuspensioncould greatly influence the food web organization through thequantity, quality and size of edible particles available ata given time.