Fecal pellets flux at DYFAMED sediment traps

Because zooplankton feces represent a potentially important transport pathway of surface-derived organic carbon in the ocean, we must understand the patterns of fecal pellet abundance and carbon mobilization over a variety of spatial and temporal scales. To assess depth-specific water column variations of fecal pellets on a seasonal scale, vertical fluxes of zooplankton fecal pellets were quantified and their contribution to mass and particulate carbon were computed during 1990 at 200, 500, 1000, and 2000 m depths in the open northwestern Mediterranean Sea as part of the French-JGOFS DYFAMED Program. Depth-averaged daily fecal pellet flux was temporally variable, ranging from 3.04 * 10**4 pellets m**2/d in May to a low of 6.98 * 10**2 pellets m**2/d in September. The peak flux accounted for 50% of the integrated annual flux of fecal pellets and 62% of pellet carbon during only two months in mid-spring (April and May). Highest numerical fluxes were encountered at 1000 m, suggesting fecal pellet generation well below the euphotic zone. However, there was a trend toward lower pellet carbon with increasing depth, suggesting bacterial degradation or in situ repackaging as pellets sink through the water column. At 500 m, both the lowest pellet numerical abundance and carbon flux were evident during the spring peak. Combined with data indicating that numerical and carbon fluxes are dominated at 500 m by a distinct type of pellet found uniquely at this depth, these trends suggest the presence of an undescribed mid-water macro-zooplankton or micro-nekton community. Fecal pellet carbon flux was highest at 200 m and varied with depth independently of overall particulate carbon, which was greatest at 500 m. Morphologically distinct types of pellets dominated the numerical and carbon fluxes. Small elliptical and spherical pellets accounted for 88% of the numerical flux, while larger cylindrical pellets, although relatively rare (<10%), accounted for almost 40% of the overall pellet carbon flux. Cylindrical pellets dominated the pellet carbon flux at all depths except 500 m, where a large subtype of elliptical pellet, found only at that depth, was responsible for the majority of pellet carbon flux. Overall during 1990, fecal pellets were responsible for a depth-integrated annual average flux of 1.03 mgC/m**2/d, representing 18% of the total carbon flux. The proportion of vertical carbon flux attributed to fecal pellets varied from 3 to 35%, with higher values occurring during periods when the water column was vertically mixed. Especially during these times, fecal pellets are a critical conveyor of carbon to the deep sea in this region.

Biological traits, and egg and fecal pellet production of Calanus finmarchicus and Calanus glacialis in situ and during experiments

The effects of temperature and food was examined for Calanus finmarchicus and C. glacialis during 3 phases of the phytoplankton spring bloom in Disko Bay, western Greenland. The 2 species were collected during pre-bloom, bloom, and post-bloom and exposed to temperatures from 0 to 10°C, combined with deficient or excess food. Fecal pellet and egg production were measured as indices for grazing and secondary production, respectively. Furthermore, changes in body carbon, nitrogen, and lipid content were measured. C. glacialis sampled before the bloom and incubated with excess food exhibited high specific egg production at temperatures between 0 and 2.5°C. Higher temperatures did not increase egg production considerably, whereas egg production for C. finmarchicus more than tripled between 2.5 and 5°C. Starved C. glacialis produced eggs at all temperatures stimulated by increasing temperatures, whereas starved C. finmarchicus needed temperatures above 5°C to produce eggs fueled by their lipid stores. Few C. finmarchicus had mature gonads at the initiation of the pre-bloom and bloom experiment, and egg production of C. finmarchicus therefore only increased as the ratio of individuals with mature gonads increased. During the bloom, both C. glacialis and C. finmarchicus used the high food availability for egg production, while refueling or exhausting their lipid stores, respectively. Finally, during the post-bloom experiment, production was low by C. finmarchicus, whereas C. glacialis had terminated production. Our results suggest that a future warmer ocean will reduce the advantage of early spawning by C. glacialis and that C. finmarchicus will become increasingly prevalent.

Production, oxygen uptake, and sinking velocity of copepod fecal pellets originated from the central North Sea

Production, oxygen uptake, and sinking velocity of copepod fecal pellets egested by Temora longicornis were measured using a nanoflagellate (Rhodomonas sp.), a diatom (Thalassiosira weissflogii), or a coccolithophorid (Emiliania huxleyi) as food sources. Fecal pellet production varied between 0.8 pellets ind**-1 h**-1 and 3.8 pellets ind**-1 h**-1 and was significantly higher with T. weissflogii than with the other food sources. Average pellet size varied between 2.2 x 10**5 µm**3 and 10.0 x 10**5 µm**3. Using an oxygen microsensor, small-scale oxygen fluxes and microbial respiration rates were measured directly with a spatial resolution of 2 µm at the interface of copepod fecal pellets and the surrounding water. Averaged volume-specific respiration rates were 4.12 fmol O2 µm**-3 d**-1, 2.86 fmol O2 µm**-3 d**-1, and 0.73 fmol O2 µm**-3 d**-1 in pellets produced on Rhodomonas sp., T. weissflogii, and E. huxleyi, respectively. The average carbon-specific respiration rate was 0.15 d**-1 independent on diet (range: 0.08-0.21 d**-1). Because of ballasting of opal and calcite, sinking velocities were significantly higher for pellets produced on T. weissflogii (322 +- 169 m d**-1) and E. huxleyi (200 +- 93 m d**-1) than on Rhodomonas sp. (35 +- 29 m d**-1). Preservation of carbon was estimated to be approximately 10-fold higher in fecal pellets produced when T. longicornis was fed E. huxleyi or T. weissflogii rather than Rhodomonas sp. Our study directly demonstrates that ballast increases the sinking rate of freshly produced copepod fecal pellets but does not protect them from decomposition.

Copepod abundance and fecal pellet production, sinking rate, and degradation in samples obtained during Leg 7 of Galathea 3 cruise 2006

The vertical distribution of copepods, fecal pellets and the fecal pellet production of copepods were measured at seven stations across the Southern Indian Ocean from productive areas off South Africa to oligotrophic waters off Northern Australia during October/November 2006. We quantified export of copepod fecal pellet from surface waters and how much was retained. Furthermore, the potential impact of Oncaea spp. and harpacticoid copepods on fecal pellets degradation was evaluated and found to be regional substantial. The highest copepod abundance and fecal pellet production was found in the western nutrient-rich stations close to South Africa and the lowest at the central oligotrophic stations. The in situ copepod fecal pellet production varied between 1 and 1,000 µg C/m**3/day. At all stations, the retention of fecal pellets in the upper 400 m of the water column was more than 99% and the vertical export of fecal pellets was low (<0.02 mg/m**2/day).

Sampling data, experimental data and fecal pellet characteristics of water pump samples in the strait of Øresund between Denmark and Sweden during 2004/2005

Copepod fecal pellets are often degraded at high rates within the upper part of the water column. However, the identity of the degraders and the processes governing the degradation remain unresolved. To identify the pellet degraders we collected water from Øresund (Denmark) approximately every second month from July 2004 to July 2005. These water samples were divided into 5 fractions (<0.2, <2, <20, <100, <200 µm) and total (unfractionated). We determined fecal pellet degradation rate and species composition of the plankton from triplicate incubations of each fraction and a known, added amount of fecal pellets. The total degradation rate of pellets by the natural plankton community of Øresund followed the phytoplankton biomass, with maximum degradation rate during the spring bloom (2.5 ± 0.49 d**-1) and minimum (0.52 ± 0.14 d**-1) during late winter. Total pellet removal rate ranged from 22% d**-1 (July 2005) to 87% d**-1 (May). Protozooplankton (dinoflagellates and ciliates) in the size range of 20 to 100 µm were the key degraders of the fecal pellets, contributing from 15 to 53% of the total degradation rate. Free-living in situ bacteria did not affect pellet degradation rate significantly; however, culture-originating bacteria introduced in association with the pellets contributed up to 59% of the total degradation rate. An effect of late-stage copepod nauplii (>200 µm) was indicated, but this was not a dominating degradation process. Mesozooplankton did not contribute significantly to the degradation. However, grazing of mesozooplankton on the pellet degraders impacts pellet degradation rate indirectly. In conclusion, protozooplankton seems to include the key organisms for the recycling of copepod fecal pellets in the water column, both through the microbial loop and, especially, by functioning as an effective 'protozoan filter' for fecal pellets.

Fecal pellet ingestion rate and feeding behavior of Oithona similis, Calanus helgolandicus and Pseudocalanus elongatus from the North Sea

Studies of fecal pellet flux show that a large percentage of pellets produced in the upper ocean is degraded within the surface waters. It is therefore important to investigate these degradation mechanisms to understand the role of fecal pellets in the oceanic carbon cycle. Degradation of pellets is mainly thought to be caused by coprophagy (ingestion of fecal pellets) by copepods, and especially by the ubiquitous copepods Oithona spp. We examined fecal pellet ingestion rate and feeding behavior of O. similis and 2 other dominant copepod species from the North Sea (Calanus helgolandicus and Pseudocalanus elongatus). All investigations were done with fecal pellets as the sole food source and with fecal pellets offered together with an alternative suitable food source. The ingestion of fecal pellets by all 3 copepod species was highest when offered together with an alternative food source. No feeding behavior was determined for O. similis due to the lack of pellet capture in those experiments. Fecal pellets offered together with an alternative food source increased the filtration activity by C. helgolandicus and P. elongatus and thereby the number of pellets caught in their feeding current. However, most pellets were rejected immediately after capture and were often fragmented during rejection. Actual ingestion of captured pellets was rare (<37% for C. helgolandicus and <24% for P. elongatus), and only small pellet fragments were ingested unintentionally along with alternative food. We therefore suggest coprorhexy (fragmentation of pellets) to be the main effect of copepods on the vertical flux of fecal pellets. Coprorhexy turns the pellets into smaller, slower-sinking particles that can then be degraded by other organisms such as bacteria and protozooplankton.

Fecal pellet and egg production rates of copepod species at varying suspended sediment concentrations

We investigated the effect of suspended sediments on the vital rates of the copepods Calanus finmarchicus, Pseudocalanus sp. and Metridia longa in a Greenland sub-Arctic fjord. The fjord had a gradient of suspended particulate matter (SPM) with high concentrations (>50 mg/L) in the inner fjord due to glacial melt water runoff. Laboratory experiments showed that when feeding on the diatom Thalassiosira weissflogii specific ingestion rates were low at high concentrations of suspended sediment for C. finmarchicus (>20 mg/L) and Pseudocalanus sp. (>50 mg/L), while no effect was found for M. longa. For C. finmarchicus, a relatively constant fecal pellet production (FPP) and fecal pellet volume suggested ingestion of sediment, which probably led to reduction in egg production rates (EPRs) at high sediment concentrations. For Pseudocalanus sp., FPP decreased with increasing sediment concentrations, while no effect was observed on EPR. No significant difference was observed in FPP for M. longa feeding on the diatom T. weissflogii compared to the ciliate Strombidium sulcatum. The study shows that high sediment concentrations influence the capability of carbon turnover in C. finmarchicus and Pseudocalanus sp., while M. longa appears to be more tolerant to high sediment loads. Therefore, high concentrations of SPM could potentially influence the species composition of glacially influenced fjords.