Profiles of photosynthetic pigments in the ocean using thin-layer chromatography

Phytoplankton pigments at a coastal station off Sydney, Australia, were studied by cellulose thinlayer chromatography. The chromatographic procedure distinguished chlorophylls a, b and c, chlorophyllide a, pheophytin a and pheophorbide a, and the major carotenoids carotene, astaxanthin, fucoxanthin, peridinin, diadinoxanthin and neoxanthin. Chlorophyllide a and pheophorbide a were the most significant chlorophyll-a degradation products in the water column, chlorophyllide a coming from chlorophyllase activity of senescent diatoms, and pheophorbide a from faecal pellets of copepods. Chlorophyll c occurred in every sample, even where there was no trace of chlorophyll a. Because of the greater chemical and biological stability of chlorophyll c compared to chlorophyll a, high chlorophyll c:a ratios result from a large proportion of senescent or detrital material in the samples. Determining the position of patches of copepods, diatoms, green algae and dinoflagellates in the water column was easily done by noting the presence of definitive pigments on the chromatograms.     

Suspended particulates in the Irish Sea and feeding conditions for fish larvae

The size distributions (2 to 160 m equivalent spherical diameter) of suspended particulate material sampled on two cruises along a transect in the Irish Sea in 1988 are described in relation to hydrographic conditions, chlorophylla concentration and carbon to nitrogen ratios. Particulates were more abundant and larger size modes in the distribution were more evident, in the upper mixed layer of stratified water than in areas where the water column was fully mixed. The detrital content was estimated at 52% of total particulate matter above the thermocline in stratified regions and at around 97% at mixed water sites. In stratified regions the predominance of larger sized phytoplankton and lower levels of detritus is argued to support a more direct and efficient transfer of energy to fish larvae via larger sizes of copepods. Conversely, in mixed areas of high detrital loading the smaller size spectrum of particulates incorporates a less efficient transfer of energy through bacterial cycling and smaller copepods.     

Effects of detrital non-native and native macroalgae on the nitrogen and carbon cycling in intertidal sediments

The decay of non-native and native seaweed mixing may modify sediment biogeochemistry and organic matter transfers within benthic food webs according to their composition and biomass. The non-native species Sargassum muticum was deliberately added to the sediment of an intertidal sandflat at different biomass and mixed to the native species Ulva sp. and Fucus vesiculosus. The sediment porewater was then ¹³C and ¹⁵N enriched to test whether both detrital diversity and biomass influenced the transfer of porewater carbon and nitrogen to the sediment and to the macrofauna consumers. More ¹⁵N-nitrogen was mobilized to sediments and macrofauna when the 3-species detrital mixing was buried, probably because this mixing provided species-specific compounds such as polyphenols due to the presence of S. muticum and F. vesiculosus, as well as large amounts of nitrogen due to the presence of Ulva. Our study revealed the importance of detrital diversity and non-native seaweeds for the nitrogen cycling in the benthic food web.     

Fate of domoic acid ingested by the copepod <Emphasis Type="Italic">Acartia clausi</Emphasis>

Two important issues in the studies of harmful algae include ecological role of the toxic compounds and their fate through the food web. The aims of this study were to determine whether the production of domoic acid is a strategy evolved to avoid predation and the role of copepods in the fate of this toxic compound through the food web. The copepod Acartia clausi was fed with single and mixed cultures of the toxic diatom Pseudo-nitzschia multiseries and the non-toxic diatom Pseudo-nitzschia delicatissima. Ingestion rate as a function of diatom abundance was the same for the toxic and non-toxic Pseudo-nitzschia species, indicating no selective feeding behaviour against P. multiseries. The toxins ingested by the copepods did not affect mortality, feeding behaviour, egg production and egg hatching of the copepods. Copepods assimilated the 4.8% of the total domoic acid ingested. Although the amount of toxins daily detoxificated by the copepods was 63.6%, the copepods accumulated domoic acid in their tissues. We conclude that domoic acid is not toxic for copepods and, probably for this reason, this toxin does not act as feeding deterrent for copepods. However, even though the production of domoic acid has apparently not evolved to deter predation, copepods may play an important role on the fate of this toxic compound through the marine food web.     

Regulation of body dimethylsulfoniopropionate (DMSP) content by the copepod Temora longicornis: a test of four mechanisms

Copepods contain dimethylsulfoniopropionate (DMSP) in their tissues in addition to the DMSP in their gut contents and therefore constitute an additional pool of DMSP in the ocean. In the estuarine copepod Temora longicornis Müller, this dynamic pool of DMSP is reduced when external salinity decreases. In the present study the mechanism(s) used by T. longicornis to adjust its DMSP content were examined. Four possible mechanisms were tested in experiments conducted between July 1997 and May 1999: (1) DMSP cleavage, (2) demethylation, (3) conversion to methionine, and (4) release from the body. Tissue extracts of T. longicornis did not exhibit the ability to cleave or demethylate DMSP. In incubation experiments, when external salinity decreased, T. longicornis individuals reduced their DMSP content without increasing their methionine content. The total amount of DMSP in the incubation vials was conserved regardless of salinity. The copepods retained most of the DMSP in their tissues in 30‰ water, but when salinity was reduced to 20‰, the copepods released 41 to 55% of their body DMSP to the surrounding medium; 89 to 91% of the total DMSP was recovered. This suggests that estuarine copepods represent a reservoir of DMSP, which is released as dissolved DMSP upon salinity decline. Therefore, while osmoregulation by these copepods may have little impact on the chemical properties of DMSP, it will affect the partitioning of DMSP between particulate and dissolved phases in the water column. Received: 20 August 1999 / Accepted: 28 March 2000     

Short-term effects of sucralose on Calanus finmarchicus and Calanus glacialis in Disko Bay,Greenland

The potential effects of sucralose on the Arctic copepods Calanus finmarchicus and Calanus glacialis were studied in Disko Bay, Greenland. Sucralose is a non-calorie sweetener and chlorine derivate of sucrose containing three chlorine atoms. Scandinavian screening studies of sucralose in 2007, revealed sucralose in all effluent samples. To investigate whether sucralose is harmful to the Arctic aquatic ecosystems, possible short-term effects were investigated on egg production, hatching rate, food intake and mortality of two species of Arctic copepods. The copepods were exposed to six different concentrations (0–50,000 ng · L^?1) of sucralose, which spans the range of concentrations found in the screening studies. Exposure led to no mortality among the copepods. Food intake by C. glacialis increased with increasing concentrations of sucralose. In C. finmarchicus, food intake did not differ with increasing concentrations. No effect of sucralose was observed on egg production of C. finmarchicus. Despite increased food intake with increasing concentrations of sucralose, C. glacialis did not increase its egg production. The results show that both species responded weakly to sucralose, but with C. glacialis being possibly slightly more sensitive to sucralose than C. finmarchicus.     

Acute physiological reactions of Calanus pacificus to selected dinoflagellates: direct observations

Calanus pacificus (Copepoda: Calanoida) females were collected off the California (USA) coast from November 1984–April 1985. A video system was used to observe and record the behavior of restrained individual females presented with a variety of dinoflagellate prey. Two species, Gonyaulax grindleyi and Ptychodiscus brevis, elicited acute physiological reactions. In 40% of the trials (n=10), copepods fed G. grindleyi regurgitated after 45 to 120 min and, in nearly all cases, did not maintain full guts. Copepods in the presence of P. brevis exhibited rapid heart rate and loss of motor control. Scrippsiella trochoidea elicited an intermediate response by C. pacificus. The copepods occasionally displayed mouthpart twitching or failure to maintain gut fullness. Olisthodiscus luteus elicited no unusual behavior in an intermediate temporal range (sec-hours), although the mouthpart movements appeared different than in copepods fed Gyrodinium resplendens (used as control). Placing the copepods in G. resplendens suspension restored normal feeding behavior in all cases.     

Biosynthesis of trimethylamine oxide in calanoid copepods. seasonal changes in trimethylamine monooxygenase activity

Live copepods, Calanus finmarchicus (Gunnerus) and C. hyperboreus (Krøyer), exposed to dissolved ¹⁴C-labeled trimethylamine (TMA) in seawater, oxidized TMA to trimethylamine oxide (TMAO), which accumulated in the organisms. The amount of TMAO synthesized was dependent on the time of exposure to TMA and the concentration of TMA in the seawater. It was inferred that copepods can produce TMAO by oxidation of TMA found in their plant food. Choline and methionine did not appear to be of importance as precursors of TMAO. There were large seasonal changes in TMA monooxygenase activity of both copepods. The activity was high in spring, and decreased through summer and autumn to a winter low in October to March. The changes in TMAO content of C. finmarchicus were, in comparison, small (this aspect was not tested for C. hyperboreus).     

A commensal relationship between the scyphozoan medusae<Emphasis Type="Italic"> Catostylus mosaicus</Emphasis> and the copepod<Emphasis Type="Italic"> Paramacrochiron maximum</Emphasis>

The copepod Paramacrochiron maximum was found in high numbers (up to 5,675 copepods/medusa) on the oral arms of the scyphozoan Catostylus mosaicus. This association was considered to be commensalism for the following reasons: P. maximum (Lichomolgidae) was abundant on the medusae (approximately 805 copepods/kg of medusae) and very rare in the water column (approximately 5.99×10^-4 copepods/kg of water); copepodites and adults of the symbiont were present on the host; the copepods were on the medusae both day and night, at different times (nine occasions between March 1999 and May 2000) and different locations (Botany Bay and Lake Illawarra, NSW, Australia). Over 40 taxa of plankton were found on the oral arms of C. mosaicus (including protists, cnidarians, polychaetes, molluscs, a wide range of holoplanktonic and meroplanktonic crustaceans, chaetognaths and fish eggs). These taxa were abundant in the water column and we concluded that they were prey. Symbiotic amphipods and carangid fishes were found with medusae. We conclude that there is a symbiotic association between P. maximum and C. mosaicus and care should be taken not to confound these copepods with the prey of C. mosaicus. Poecilostomid copepods are well known for consuming mucus and feeding is likely to be a major reason for the association.Communicated by G.F. Humphrey, Sydney     

A continuous recirculating culture system for planktonic copepods

The calanoid copepods, Acartia clausi Giesbrecht and Acartia tonsa Dana, are maintained at high densities in continuous culture at 15°C. Synthetic sea-water medium is recirculated through filters and a foam tower which limits accumulation of dissolved wastes and various metabolites. The ciliate Euplotes vannus Müller is associated in culture with the copepods, and effectively controls bacterial population and accumulation of algal debris. The copepods graze upon the ciliates as well as upon the phytoflagellates Isochrysis galbana Parke and Rhodomonas baltica Korsten.Contribution No. 119 from the Institute of Marine and Atmospheric Sciences, University of Miami.     

Vertical distribution and nocturnal migration of zooplankton in relation to the development of the seasonal thermocline in Patraikos Gulf

Vertical distribution and nocturnal migration of zooplankton species in relation to the development of the seasonal thermocline in the shallow waters (90 m) of Patraikos Gulf (Ionian Sea, Greece) were investigated using a WP-2 closing net. Juvenile and adult copepods accounted for a mean of 91% of the total collected in three sampling periods, i.e. May, July and September 1985.Ctenocalananus vanus, Paracalanus parvus andOithona plumifera were the dominant copepods. The majority of the zooplankton tend to aggregate at the thermocline layer. Among copepods the two congeneric speciesClausocalanus pergens andC. furcatus exhibited different migratory responses to the development of the thermocline.C. pergens occurred in the lower part of the thermocline andC. furcatus in the upper region or above. The diel vertical migration of all species could be divided into four types: (1) no vertical migration; (2) upward migration at night; (3) occasional migration; and (4) reverse migration (down at night). In July when the strongest thermocline developed, most zooplankters rose close to the surface at night. For most species, temperature discontinuity did not limit their diel migration.Please address all correspondence and requests for reprints to Dr J.J. Lykakis     

Feeding four species of pelagic copepods under experimental conditions

Different qualitative and quantitative aspects of feeding with varied compositions of diets have been studied in 4 species of pelagic copepods: Calanus helgolandicus, Centropages typicus, Temora stylifera and Acartia clausi. By feeding copepods different algal concentrations, it was shown that when food concentration increases grazing rate decreases; the ingestion rate remains fairly constant in the lower range of concentrations, but then increases, reaching a plateau at higher algal concentrations. There is a significant correlation between daily food intake and fecal pellet production. On a pluri-algal diet, selective grazing is observed: larger phytoplankton cells are more efficiently removed than smaller ones. Using Artemia nauplii, it is shown that the copepods studied are also able to eatch and ingest animal prey. Increased daily food intake affects respiration and oviposition. Metabolic requirements, gross growth-efficiency, and food assimilation have been ealculated in Calanus helgolandicus, Centropages typicus and T. stylifera for a large range of algal concentrations.     

Impact of copepod grazing on the red-tide flagellate Chattonella antiqua

Although planktonic copepods are major suspension feeders in the sea, the impact of their grazing pressure upon red-tide flagellates has not been fully investigated. In the present study, the grazing of adult females of several copepod species is examined using three food types: viz. natural suspended particles, natural suspended particles mixed with cultured Chattonella antiqua, and cultured C. antiqua. The functional response on C. antiqua was investigated for five species of copepods (Acartia erythraea, Calanus sinicus, Centropages yamadai, Paracalanus parvus and Pseudodiaptomus marinus). Ingestion rates increased linearly with increasing cell concentrations until a maximum level was reached, beyond which the rates were constant. This cell concentration was higher for larger copepods. The weight-specific maximum ingestion rates were higher in the small species. In general, copepods tended to feed selectively on larger particles when feeding on natural particles. This tendency was strongest in a simulated red-tide environment. Thus, it can be surmised that copepods may selectively graze on C. antiqua during the outbreak of a red tide. Grazing pressure by the natural copepod community in Harima Nada, the Inland Sea of Japan, was calculated by integration of the laboratory determined feeding rates and field measurements of zooplankton biomass. The daily removal rate was 3.4 to 30.8% (mean: 12.3%) of C. antiqua biomass at 20 cells ml^-1 and decreased to 0.6–4.3% (mean: 1.8%) at 500 cells ml^-1. Therefore, the grazing pressure by the copepod community is important at the initial stage of the red tide.     

Diet of copepods (Scopalatum vorax) associated with mesopelagic detritus (giant larvacean houses) in Monterey Bay,California

The feeding structures or houses of the giant larvacean Bathochordaeus sp. serve as both habitat and food for the calanoid copepod Scopalatum vorax. Gut contents of S. vorax include both microbial and metazoan associates of larvacean houses, and possibly the house-mucus matrix itself. Copepods were observed and collected from larvacean houses between 100 and 500 m in Monterey Bay, California, using a submersible ROV (remotely operated vehicle) from the Monterey Bay Aquarium Research Institute. Gut contents were compared to potential food items on the houses and in the open water (not associated with the house). Copepods were generalist feeders, with amorphous detritus, diatoms, and copepods or other crustacean parts dominating gut contents. Protozoans and algae other than diatoms were rarer in guts. Houses contained a diverse assemblage of microplankton and metazoans, both intact specimens and detrital remains of these. Numbers of diatoms and fecal pellets were enriched by 1 to 3 orders of magnitude on houses compared to numbers in surrounding water. Many of the abundant species of diatoms and copepods on houses occurred in S. vorax guts. This observation coupled with S. vorax feeding habits observed in situ and in the laboratory provide evidence for feeding on houses. S. vorax appears to possess special adaptations to living in a resource-limited environment, such as gorging as a feeding adaptation, chemosensory structures to help locate houses, and the ability to change feeding modes. Consumption of detritus at depth by S. vorax provides evidence that metazoans contribute to remineralization of particulate organic carbon in the mesopelagic zone.     

Omnivorous feeding behavior of the Antarctic krill Euphausia superba

Feeding experiments were conducted at Palmer Station from December 1985 to February 1986 to examine the potential role of copepod prey as an alternative food source for Euphausia superba. Copepod concentration, copepod size, phytoplankton concentration, the duration of krill starvation and the volume of experimental vessels were altered to determine effects on ingestion and clearance rates. Krill allowed to feed on phytoplankton and copepods in 50-litre tubs showed greatly increased feeding rates relative to animals feeding in the much smaller volumes of water traditionally used for krill-feeding studies. Clearance rates on copepods remained constant over the range of concentrations offered, but clearance rates on phytoplankton increased linearly with phytoplankton concentration. Feeding rates increased when larger copepods were offered and when krill were starved for two weeks prior to experiments. Clearance rates of krill feeding on copepods were higher than, but not correlated with, their clearance rates on phytoplankton in the same vessel. E. superba may have a distinct mechanism for capturing copepods, perhaps through mechanoreception. Although our observed clearance rate of 1055 ml krill^-1 h^-1 indicates that krill can feed very efficiently on copepod prey, such feeding would meet less than 10% of their minimum metabolic requirements at the typical copepod concentrations reported for Antarctic waters. However, substantial energy could be gained if krill fed on the patches of high copepod concentrations occasionally reported during the austral summer, or if krill and copepods were concentrated beneath the sea ice during the winter or spring months. Our results, indicating efficient feeding on zooplankton and higher clearance rates on phytoplankton than previously believed, represent a step towards balancing the energy budget of E. superba in Antarctic waters.     

Differing distributions of potentially competing amphipods,copepods and gastropods among specimens of the intertidal alga Pelvetia fastigiata

Possible competitive interactions are inferred from the distributions and abundances of species within three invertebrate groups (Hyale spp., Ampithoe spp. and thallusdwelling species) among unmanipulated and experimental specimens of the brown alga Pelvetia fastigiata collected between 1974 and 1976, inclusive, at La Jolla, California, USA. During the fall, when the vagile amphipods Hyale grandicornis and H. frequens were relatively abundant, they were found mostly on specimens of P. fastigiata at upper mid-intertidal and mid-tide levels, respectively. H. grandicornis often dispersed to mid-tide levels, but the two species cooccurred on few plants. Although H. frequens infrequently occupied P. fastigiata in the spring, H. grandicornis still generally sheltered in upper mid-intertidal plants, in part, because the net reproduction rates of H. grandicornis were low when space was available. The tubiculous amphipod A. tea usually was the only Ampithoe species on P. fastigiata; however, A. lindbergii and A. pollex sometimes dispersed to and released broods on this alga, particularly during the early summer when abundances of A. tea were relatively low. Similarly, net immigration rates of other thallus-dwelling copepods were relatively high when abundances of Scutellidium lamellipes were low. Moreover, several copepods were most abundant on different specimens of small isolated plants, and thallus-dwelling gastropods on larger plants at sites with relatively low densities of S. lamellipes on P. fastigiata; such copepods and gastropods generally were not abundant on aggregated plants sheltering large numbers of S. lamellipes. Such differences suggest that competition for food or space is among the factors causing negative correlations in the densities of Ampithoe spp. and thallusdwelling species on P. fastigiata.     

Comparison of the food of Triphoturus mexicanus and T. nigrescens,two lanternfishes of the Pacific Ocean

Prey (chiefly euphausiids and copepods) eaten by two myctophids (lanternfishes) are compared from incidence in fish stomachs and from abundance in the environment. One lanternfish species, Triphoturus mexicanus, lives in the California Current, and the other, T. nigrescens, lives in the central Pacific Ocean. Although these two environments are very different physically and biologically, the feeding habits of the two lanternfishes are surprisingly similar. Prey biomass is 94% euphausiids, 3% copepods, and 3% other organisms for T. mexicanus and 88% euphausiids, 4.5% copepods, and 7.5% other organisms for T. nigrescens; the difference between the fish species is not significant when tested statistically. The two fishes resemble one another in frequency distributions of ingested copepod individuals, copepod species, euphausiid individuals, and euphausiid species. During a single diurnal feeding period, both fishes eat a variety of copepod species but tend to eat only a single species of euphausiid. T. mexicanus grows to twice the length of T. nigrescens and eats proportionally larger euphausiids; however, both fishes eat copepods having the same median size. The frequencies of euphausiid species in the diets of both fishes differ from the frequencies in the environment. The chief differences between the feeding habits of the two lanternfishes are that T. nigrescens, in comparison to its congener, eats a greater variety of organisms during one diurnal feeding period and captures smaller euphausiids. The feeding patterns for each lanternfish species are consistent over distances of hundreds of kilometers and over many years of sampling.     

Plankton community dynamics of the central Great Barrier Reef Lagoon: Analysis of data from Ikeda et al.

Plankton data collected by Ikeda et al. (1980) from the central region of the Great Barrier Reef, and spanning two years (1976 through 1978) of zooplankton records, have been analyzed extensively for spatial and temporal patterns. Estimates of net zooplankton (including chaetognaths, copepods, and larvaceans) and microzooplankton (juvenile copepods, encompassing nauplii and copepodites, and ciliates) were assessed at three stations across the 60 km lagoon. Temperature, salinity, and chlorophyll a were also measured. A cross-lagoonal gradient was identified in the plankton, concurring with results of related surveys of benthic taxa, such as scleractinian corals, soft corals, macro-algae, fish, sponges, crinoids, etc. Two associations of net zooplankton were identified. The first was associated primarily with the inner lagoon; the second with the outer lagoon. The inshore association was characterized by higher abundances of almost all net zooplankton taxa, particularly chaetognaths, copepods, polychaetes, decapods, and meroplanktonic larvae as well as higher concentrations of chlorophyll a. This inshore association wove back and forth across the lagoon through time, dominating the lagoon entirely during periods of high river discharge, reaching the mid-shelf platform reefs in this region, and sometimes being entirely absent during dry periods. Both seasonal and annual peaks in plankton abundance were generally linked with degree of runoff. Summer/autumn peaks of abundance were evident in chaetognaths, copepods, and larvaceans while annual variation was detected in the former two as well as in chlorophyll a concentrations. Depth stratification was noted in juvenile copepods and chlorophyll a concentrations at the center of the lagoon, with higher abundances recorded in deeper waters. The central Great Barrier Reef lagoon was found to be typical of other tropical coastal waters where plankton community dynamics are controlled primarily by physical factors. We suggest that any substantial changes in river discharge in this area will affect plankton production.A.I.M.S. Contribution No. 242     

Active swimming in meiobenthic copepods of seagrass beds: geographic comparisons of abundances and reproductive characteristics

The entry of meiobenthic copepods from sediments or seagrass blades into the water column and reproductive characteristics of actively migrating fauna were investigated from 1981–1986 in a temperate intertidal Zostera capricorni seagrass bed in Pautahanui Inlet, New Zealand and in a subtidal Thalassia testudinum bed in Tampa Bay, Florida, USA. Emergence of copepods in New Zealand varied over a tidalcycle, while in Florida a distinct diel periodicity was displayed. Selected copepod species in New Zealand had similar numbers emerging from sediments and/or blades over a 6 h period as the common copepods actively migrating from sediments in Florida. Daily abundances of emerging copepods (24 h) in Tampa Bay, Florida, were substantially greater than those in New Zealand, where migration is linked to tidal cover. In Z. capricorni meadows in New Zealand, sex ratios of copepods in sediments and on blades were dominated by females; males dominated water-column samples. In T. testudinum meadows in Tampa Bay, sex ratios of males to females, although of a lowermagnitude than in Z. capricorni beds, were higher in trap than in sediment samples. Differences in sex ratios, the availability in emergence traps of females of appropriate stage for mating, and observations on clasping in live samples from traps suggest that swimming behavior in copepods may be partly linked to prenuptial courtship. Meiobenthic copepods may use the water column as an important habitat for reproductive behavior.     

Increased production of faecal pellets by the benthic harpacticoid Paramphiascella fulvofasciata: importance of the food source

The re-use of faecal pellets in the water column before sinking to the seafloor is known as an important pathway in marine food webs. Especially planktonic copepods seems to be actively use their faecal pellets. Since benthic copepods (order Harpacticoida) live in the vicinity of their pellets, it remains unclear how important these pellets are for their feeding ecology. In the present study a laboratory experiment was conducted to analyse the importance of faecal pellets for the feeding ecology of the harpacticoid Paramphiascella fulvofasciata and its grazing pressure on two diatom species (Seminavis robusta, Navicula phyllepta). By quantifying the amount and volume of the produced faecel pellets in different treatments, it was tested to what extent the food source and the lack of faecal pellets would influence the production of faecal pellets. We found that the grazing pressure of P. fulvofasciata depended on the diatom density since only a top-down effect could be found on the smaller Navicula cells during its initial exponential growth phase. The grazer had a negative effect on the diatom growth and controlled the cell density to about 4,000 cells/cm². In spite of the fact that the addition of faecal pellets did not show a significant positive effect on the assimilation of diatoms, the removal of faecal pellets strongly promoted the pellet production. Especially when grazing on Navicula the harpacticoid P. fulvofasciata produced significantly more and smaller faecal pellets when the pellets were removed. This outcome illustrates the need for faecal pellets of this harpacticoid copepod when grazing on the diatom Navicula. Apart from its selection for smaller diatom cells, it was suggested that the colonisation of heterotrophic bacteria enriched these pellets. This study is the first to indicate that trophic upgrading occurs on faecal pellets and not only on the initial autotrophic food sources per se.