Gut clearance and pigment destruction in a herbivorous copepod, Acartia tonsa, and the determination of in situ grazing rates

Gut clearance rates of starving and continuously feeding Acartiatonsa were estimated. During the initial 30 min the rates weresimilar (0.045 and 0.048 min^–1, respectively; 14°C)but thereafter starving animals expelled the remains of theirgut contents at half the rate (0.019 min^–1) of fed ones(0.048). Pigment destruction was estimated by (i) incubationexperiments over 3–4 days, (ii) silica to pigment ratioin algae and faeces and (iii) by gut filling experiments. Theincubations showed that 8% of the ingested pigments were destroyedto nonfluorescent residues during gut passage. The silica topigment ratio method gave an average of 11 % (1 –24) destructionand gut-filling experiments showed no systematic differencebetween ingestion measured as gut filling rate (fluorescence)and particle reduction. ¹Present address: Kristineberg Marine Biological Station, S-45034 Fiskebäckskil, Sweden     

The Sea Core Sampler: a simple water sampler that allows direct observations of undisturbed plankton

A simple water sampler that brings undisturbed water samplesto the deck-laboratory and that allows video recording of planktonand small-scale plankton phenomena directly in the sampler isdescribed. Using the sampler, it is possible to follow individualplankton organisms and marine snow aggregates for longer periodsof time in an environment where the original three-dimensionalorganization of particles and chemical gradients have been retained.The sampler has been used to film sinking marine snow, the microbiallife on a snow flake, and the behaviour of delicate zooplanktersthat are otherwise difficult to collect live. Video clips demonstratingthe utility of the sampler are available online.     

Effects of a storm event on the structure of the pelagic food web with special emphasis on planktonic ciliates

Storm-induced changes in the water column structure and thepelagic food web were investigated by daily sampling duringa 3 week period (October 24 to November 10, 1988) at a permanentstation in the southern Kattegat (Denmark) Subsequent to a stormthe primary production increased and the size-distribution ofthe phytoplankton changed towards larger cells. Some componentsof the zooplankton community responded to the increase in potentialfood, either in terms of a functional response (copepods) orin terms of a numerical response (cladocera), whereas no changesin the populations of mixo- and heterotrophic aliates were evident.The dilates were not food limited since growth rates determinedin situ were comparable to maximum growth rates measured inthe laboratory. Calculations suggest that the total clearancecapacity of planktonic copepods (as fraction of water columncleared per unit time) was of the same magnitude as the measuredinstantaneous growth rates of the ciliates (0.55–0 85day^–1). Ciliate populations were therefore probably limitedby copepod predation. Ciliates, however, contributed only insignificantlyto the diets of copepods. Although the total zooplankton productionincreased by at least 30% subsequent to the storm (and thatof the copepods by 50%), the zooplankton community was unableto assimilate the entire increase in primary production. Therelative zooplankton (copepods, cladoceran and ciliates) grazingrate on the primary production decreased 30%, and the flow ofcarbon to biomass accumulation, other grazers and in particularsedimentation is predicted to have increased by 100% followingthe storm.     

Feeding rates in the chaetognath Sagitta elegans: effects of prey size, prey swimming behaviour and small-scale turbulence

The gut contents of Sagitta elegans were sampled twice daily(noon and midnight) during 9 days in October at an anchor stationin the northern North Sea. Observations of the ambient preyfield and of turbulent dissipation rates were collected simultaneously.The average number of prey per chaetognath was among the highestever recorded, 0.57 ± 0.10. Total gut content was independentof ambient prey concentration, suggesting that feeding ratewas saturated. Clearance rates were estimated from gut contentsand ambient prey concentrations and a literature-based estimateof digestion time. The clearance rate to prey size showed adome-shaped relationship. The maximum clearance rates, about100–300 ml h^-1, were observed for prey sizes correspondingto 6–10% of Sagitta length. Clearance rates varied notonly with prey size, but also with prey type. For example, copepodmales were cleared at rates up to an order of magnitude higherthan similarly sized females, probably owing to differencesin swimming behaviour. Sagitta elegans is an ambush predatorthat perceives its prey by hydromechanical signals. Faster swimmingprey generates stronger signals and is, hence, perceived atlonger distances. We develop a simple prey encounter rate modelby describing the swimming prey as a ‘force dipole’and assuming that a critical signal strength is required toelicit an attack. By fitting the model to the observations,a critical signal strength of 10^-2 cm s^-1 is estimated; thisis very similar to estimates for copepods that also perceiveprey by mechanoreceptory setae. Gut contents were independentof turbulent dissipation rate. Because feeding rates were saturated,we did not expect to see positive effects of turbulence. However,the strong wind-generated turbulent dissipation rates observedduring the study (10^-3–10^-1 cm² s^-3 in the upper mixedlayer) could lead to negative effects by interfering with preyperception. At a dissipation rate of 10^-2 cm² s^-3 a 10-mm longS. elegans would experience fluid signals of order 0.3 cm s^-1due to turbulence, 30 times stronger than the signal strengthfrom the prey. It is, therefore, suggested that S. elegans isable to separate prey signals from turbulence signals due totheir different spatial characteristics.     

Abundance, size distribution and bacterial colonization of transparent exopolymeric particles (TEP) during spring in the Kattegat

The abundance, size distribution and bacterial colonizationof transparent exopolymeric particles (TEP) were monitored inthe Kattegat (Denmark) at weekly intervals throughout the spring(February-May) encompassing the spring diatom bloom. These recentlydiscovered particles are believed to be formed from colloidalorganic material exuded by phytoplankton and bacteria, and mayhave significant implications for pelagic flux processes. Duringthis study, the number concentration of TEP (>1 µm)ranged from 3 x 10³ to 6 x 10⁴ ml^–1 and the volume concentrationbetween 0.3 and 9.0 p.p.m.; they were most abundant in the surfacewaters subsequent to the spring phytoplankton bloom. The rangeof TEP (encased) volume concentration was similar to that ofthe phytoplankton, although at times TEP volume concentrationexceeded that of the phytoplankton by two orders of magnitude.The TEP size distribution followed a power law, with the abundanceof particles scaling with particle diameter^–(ß+1).The seasonal average estimate of ß (2.3) was not significantlydifferent from three, consistent with TEP being formed by shearcoagulation from smaller particles. However, date-specific estimatesof ß differed significantly from three, probably becauseTEP are fractal. All TEP were colonized by bacteria, and bacteriawere both attached to the surface of and embedded in TEP. Yetthe number of attached bacteria per TEP was related neitherto the surface area nor the volume, but rather scaled with TEPsize raised to an exponent of     

Effects of large gut volume in gelatinous zooplankton: ingestion rate, bolus production and food patch utilization by the jellyfish Sarsia tubulosa

Many gelatinous zooplankton consume a large amount of prey andhave stomach volumes much greater than the volume of individualprey. We suggest that jellyfish can use their voluminous stomachas a buffering food-accumulating organ that allows the organismto feed at maximum clearance rate in a wide range of fluctuatingfood concentrations. The food accumulation capability was confirmedfor the hydromedusa Sarsia tubulosa feeding on copepods. Starvedjellyfish feeding in high prey concentrations for 1 h displayedmuch higher average ingestion rates compared with jellyfishfeeding for 20 h or with jellyfish that were pre-adjusted tothe food concentration before incubation. The findings haveimplications for design and interpretation of experiments. Thepossibility for jellyfish to feed at maximum clearance ratein either very high prey concentration for a short time or lowprey concentration for a long time was illustrated with calculationsof prey uptake by S. tubulosa feeding in prey concentrationsof variable heterogeneity. The ability of jellyfish to captureprey at maximum clearance rate under different prey concentrations,and to accumulate relatively large amounts of food in theirguts, suggests that they would thrive in both homogenous andpatchy food distributions. This property may have contributedto the evolutionary and ecological success of the medusoid ‘bauplan’.     

Copepod egg production, moulting and growth rates, and secondary production, in the Skagerrak in August 1988

Measurements of hydrography, chlorophyll, moulting rates ofjuvenile copepods and egg production rates of adult female copepodswere made at eight stations along a transect across the Skagerrak.The goals of the study were to determine (i) if there were correlationsbetween spatial variations in hydrography, phytoplankton andcopepod production rates, (ii) if copepod egg production rateswere correlated with juvenile growth rates, and (iii) if therewas evidence of food-niche separation among co-occumng femalecopepods The 200 km wide Skagerrak had a stratified water columnin the center and a mixed water column along the margins. Suchspatial variations should lead to a dominance of small phytoplanktoncells in the center and large cells along the margins; however,during our study blooms of Gyrodinium aureolum and Ceratium(three species) masked any locally driven differences in cellsize: 50% of chla was >11 µm, 5% in the 11–50µm fraction and 45% <50 µm. averaged for allstations. Chlorophyll ranged from 0.2 to 2.5 µg l^–1at most depths and stations. Specific growth rates of copepodsaveraged 0.10 day^–1 for adult females and 0.27 day^–1for juveniles The latter is similar to maximum rates known fromlaboratory studies, thus were probably not food-limited. Eggproduction rates were food-limited with the degree of limitationvarying among species: 75% of maximum for Centropages typicus, 50% for Calanus finmarchicus, 30% for Paracalanus parvus and 15% for Acartia longiremis and Temora longicornis. Thedegree of limitation was unrelated to female body size suggestingfood-niche separation among adults. Copepod production, summedover all species, ranged from 3 to 8 mg carbon m^–3day^–1and averaged 4.6 mg carbon m^–1 day^–1. Egg productionaccounted for 25% of the total.     

Remote prey detection in Oithona similis: hydromechanical versus chemical cues

We quantified prey encounter rates and prey reaction distancesin the ambush-feeding cyclopoid copepod Oithona similis by videorecording freely swimming copepods at different concentrationsof prey, the dinoflagellate Gymnodinium dominans. Prey encounterrate increased with prey concentration, and a maximal clearancerate of 0.42 ± 0.10 ml h^–1 was estimated. The averagedistance (from the antennules) at which O.similis reacts toprey is 0.014 ± 0.007 cm. A simple prey encounter modelwas used to combine observed predator and prey velocities andprey reaction distance, and yielded a clearance rate similarto that estimated directly from prey encounter rates. The observedprey reaction distance was consistent with that estimated froma published model of hydromechanical prey perception. The possibilityof remote chemodetection was examined by modeling the distributionof solutes leaking out of a swimming cell. The cell leaves along slender chemical trail in its wake. However, since theambush-feeding O.similis is essentially stationary when perceivingprey, it is the width rather than the length of the trail thatmatters. Owing to advection, the chemical signal vanishes almostinstantaneously off the sides of the swimming flagellate, andsolute concentrations are below any likely detection thresholdwithin 40–50 µm from the flagellate. Our observationsare thus inconsistent with remote chemodetection in O.similis.The considerations are generalized, and it is concluded thatambush-feeding copepods, unlike cruisers and suspension feeders,cannot utilize chemical signals for the detection of individualprey, but rely on either hydromechanical detection or directinterception of prey.     

Phytoplankton aggregate formation: observations of patterns and mechanisms of cell sticking and the significance of exopolymeric material

Flocculation of ‘sticky’ phytoplankton cells intorapidly sinking aggregates has been invoked as a mechanism explainingmass sedimentation of phytoplankton blooms in the ocean. Phytoplanktonstickiness, defined as the probability of adhesion upon collision,is one key factor determining the potential for aggregate formation.In the laboratory, we examined variation in stickiness in fivespecies of diatoms and two species of flagellates grown in batchcultures. We also investigated the production of paniculatemucus by phytoplankton cells and its role in aggregate formation,and we studied the effects of solute exudates on cell stickiness.Four of the five diatoms investigated were significantly sticky,while one diatom and both of the flagellates were not sticky.Stickiness varied considerably within species. In the diatomSkeletonema costatum, the typical but not entirely consistentpattern was that stickiness decreased with age of the batchcultures. We were otherwise unable to establish consistent relationshipsbetween cell stickiness and the growth stage of the algae, environmentalconcentrations of inorganic nutrients, and abundances of suspendedand epiphytic bacteria. We showed that the diatom S.costatumat times excretes a solute substance that depresses flocculation.This may reduce cell losses from the euphotic zone during thegrowth phase due to flocculation and sedimentation. We demonstratedtwo different mechanisms of phytoplankton aggregate formation.In the diatom S.costatum, the cells are sticky in themselves,and coagulation depends on cell-cell sticking and does not involvemucus. Aggregates are composed solely of cells. Cells of thediatom Chaetoceros affinis, on the other hand, are not in themselvessticky. Transparent exopolymeric particles (TEP), produced bythe diatom, cause the cells to aggregate and coagulation dependson TEP-cell rather than cell-cell sticking. Aggregates are formedof a mixture of mucus and cells. We found several species ofdiatoms and one flagellate species to produce copious amountsof TEP. TEP from some species (e.g. Coscinodiscus sp.) is stickyand may cause other, non-sticky particles to coagulate. Thisemphasizes the potential importance of diatom-derived paniculatemucus for particle flocculation in the ocean.