Allochthonous inputs of riverine picocyanobacteria to coastal waters in the Arctic Ocean

The observed onset of climate change at high northern latitudes has highlighted the need to establish current baseline conditions in the Arctic Ocean, and has raised concern about the potential for the invasion and growth of biota that have warm temperature optima, such as cyanobacteria. In this study, we used 16S rRNA gene sequences as a molecular marker to evaluate the hypothesis that Arctic rivers provide a major inoculum of cyanobacteria into the coastal Arctic Ocean. Surface samples were collected along a transect extending from the Mackenzie River (Northwest Territories, Canada), across its estuary, to 200 km offshore at the edge of the perennial Arctic pack ice (Beaufort Sea). The highest picocyanobacteria concentrations occurred in the river, with concentrations an order of magnitude lower at offshore marine stations. The 16S rRNA gene clone libraries of five surface samples and five strains along this gradient showed that the cyanobacterial sequences were divided into eight operational taxonomic units (OTUs), six OTUs closely related to freshwater and brackish Synechococcus and two OTUs of filamentous cyanobacteria. No typically marine Synechococcus sequences and no Prochlorococcus sequences were recovered. These results are consistent with the hypothesis of an allochthonous origin of picocyanobacteria in the coastal Arctic Ocean, and imply survival but little net growth of picocyanobacteria under the present conditions in northern high-latitude seas.     

Environmental selection of marine stramenopile clades in the Arctic Ocean and coastal waters

Single-celled bacterial grazers are key components of marine ecosystems, and at times represent the dominant eukaryotes in the Arctic Ocean. Among these are small, phylogenetically diverse stramenopiles, known almost exclusively from their 18S rRNA gene sequences. Marine stramenopiles (MAST) have been found in the upper waters of the world ocean and in all Arctic seas. In particular, three sub-clades of MAST-1, MAST-1A, 1B and 1C co-occur in Arctic waters, but ecological and distributional details are lacking. Here, we have updated phylogenies of the MAST-1 clade to test whether there are Arctic-specific MAST-1 ecotypes. In addition, taking advantage of samples collected over 2 years as part of several Arctic cruises leading up to and during the International Polar Year, we applied sub-clade-specific probes and fluorescent in situ hybridization to describe the distribution of the three sub-clades in the Canadian Arctic. The three groups were found principally in the euphotic zone; however, evidence of Arctic ecotypes remains elusive. Some environment-specific separation among sub-clades was detected, with MAST-1C reaching significantly greater concentrations near the marginal ice zone, while MAST-1A and MAST-1B were associated with ice-covered and open-water stations, respectively. MAST-1B appeared to be able to persist at greater depths. Changing ice cover and mixing regimes are therefore expected to have an impact on the concentrations and distribution of these sub-clades, with possible downstream consequences on the marine food webs.     

Nitrogen metabolism by heterotrophic bacterial assemblages in Antarctic coastal waters

Field studies to examine the in situ assimilation and production of ammonium (NH₄ ⁺) by bacterial assemblages were conducted in the northern Gerlache Strait region of the Antarctic Peninsula. Short term incubations of surface waters containing ¹⁵N-NH₄ ⁺ as a tracer showed the bacterial population taking up 0.041–0.128 g-atoms Nl^–1d^–1, which was 8–25% of total NH₄ ⁺ uptake rates. The large bacterial uptake of NH₄ ⁺ occurred even at low bacterial abundance during a rich phytoplankton bloom. Estimates of bacterial production using ³H-leucine and -adenine were l.0gCl^–1 d^–1 before the bloom and 16.2 g Cl^–1 d^–1 at the bloom peak. After converting bacterial carbon production to an estimate of nitrogen demand, NH₄ ⁺ was found to supply 35–60% of bacterial nitrogen requirements. Bacterial nitrogen demand was also supported by dissolved organic nitrogen, generally in the form of amino acids. It was estimated, however, that 20–50% of the total amino acids taken up were mineralized to NH₄ ⁺. Bacterial production of NH₄ ⁺ was occurring simultaneously to its uptake and contributed 27–55% of total regenerated NH₄ ⁺ in surface waters. Using a variety of ¹⁵N-labelled amino acids it was found that the bacteria metabolized each amino acid differently. With their large mineralization of amino acids and their relatively low sinking rates, bacteria appear to be responsible for a large portion of organic matter recycling in the upper surface waters of the coastal Antarctic ecosystem.     

Rotifer distributions in the coastal waters of the northeast Pacific Ocean

Absolute abundance of rotifers was assessed from 5 to 80 km across the continental shelf off of the southern Oregon coast (U.S.A.) in the northeast Pacific Ocean. A total of 97 vertically stratified water samples were collected at 49 stations from two depths, 3 and 30 m. Coastal upwelling conditions were indicated, with decreased temperature, increased salinity and higher chlorophyll-a concentrations closer to shore. Two rotifer genera, Synchaeta and Trichocercaoccurred within 16 km of shore with densities increasing closer to shore. Synchaeta reached densities of 64 inds l^–1 while Trichocerca was sparse (<1 inds. l^–1). Rotifers were most abundant at 3 m and the densest aggregations appear to be associated with estuary outlets, suggesting that estuaries may be important in exporting rotifers to nearshore coastal waters.     

Phylogenetic composition of Arctic Ocean archaeal assemblages and comparison with Antarctic assemblages

Archaea assemblages from the Arctic Ocean and Antarctic waters were compared by PCR-denaturing gradient gel electrophoresis (DGGE) analysis of 16S rRNA genes amplified using the Archaea-specific primers 344f and 517r. Inspection of the DGGE fingerprints of 33 samples from the Arctic Ocean (from SCICEX submarine cruises in 1995, 1996, and 1997) and 7 Antarctic samples from Gerlache Strait and Dallman Bay revealed that the richness of Archaea assemblages was greater in samples from deep water than in those from the upper water column in both polar oceans. DGGE banding patterns suggested that most of the Archaea ribotypes were common to both the Arctic Ocean and the Antarctic Ocean. However, some of the Euryarchaeota ribotypes were unique to each system. Cluster analysis of DGGE fingerprints revealed no seasonal variation but supported depth-related differences in the composition of the Arctic Ocean Archaea assemblage. The phylogenetic composition of the Archaea assemblage was determined by cloning and then sequencing amplicons obtained from the Archaea-specific primers 21f and 958r. Sequences of 198 clones from nine samples covering three seasons and all depths grouped with marine group I Crenarchaeota (111 clones), marine group II Euryarchaeota (86 clones), and group IV Euryarchaeota (1 clone). A sequence obtained only from a DGGE band was similar to those of the marine group III Euryarchaeota:     

Phylogenetic Composition of Arctic Ocean Archaeal Assemblages and Comparison with Antarctic Assemblages

Archaea assemblages from the Arctic Ocean and Antarctic waters were compared by PCR-denaturing gradient gel electrophoresis (DGGE) analysis of 16S rRNA genes amplified using the Archaea-specific primers 344f and 517r. Inspection of the DGGE fingerprints of 33 samples from the Arctic Ocean (from SCICEX submarine cruises in 1995, 1996, and 1997) and 7 Antarctic samples from Gerlache Strait and Dallman Bay revealed that the richness of Archaea assemblages was greater in samples from deep water than in those from the upper water column in both polar oceans. DGGE banding patterns suggested that most of the Archaea ribotypes were common to both the Arctic Ocean and the Antarctic Ocean. However, some of the Euryarchaeota ribotypes were unique to each system. Cluster analysis of DGGE fingerprints revealed no seasonal variation but supported depth-related differences in the composition of the Arctic Ocean Archaea assemblage. The phylogenetic composition of the Archaea assemblage was determined by cloning and then sequencing amplicons obtained from the Archaea-specific primers 21f and 958r. Sequences of 198 clones from nine samples covering three seasons and all depths grouped with marine group I Crenarchaeota (111 clones), marine group II Euryarchaeota (86 clones), and group IV Euryarchaeota (1 clone). A sequence obtained only from a DGGE band was similar to those of the marine group III Euryarchaeota.     

Metabolic diversity of heterotrophic bacterioplankton over winter and spring in the coastal Arctic Ocean

Metabolic diversity of heterotrophic bacterioplankton was tracked from early winter through spring with Biolog Ecoplates under the seasonally ice covered arctic shelf in the Canadian Arctic (Franklin Bay, Beaufort Sea). Samples were taken every 6 days from December 2003 to May 2004 at the surface, the halocline where a temperature inversion occurs, and at 200 m, close to the bottom. Despite the low nutrient levels and low chlorophyll a , suggesting oligotrophy in the winter surface waters, the number of substrates used (NSU) was greater than in spring, when chlorophyll a concentrations increased. Denaturing gradient gel electrophorisis analysis also indicated that the winter and spring bacterial communities were phylogenetically distinct, with several new bands appearing in spring. In spring, the bacterial community would have access to the freshly produced organic carbon from the early phytoplankton bloom and the growth of rapidly growing specialist phenotypes would be favoured. In contrast, in winter bacterioplankton consumed more complex organic matter originated during the previous year's phytoplankton production. At the other depths we tested the NSU was similar to that for the winter surface, with no seasonal pattern. Instead, bacterioplankton metabolism seemed to be influenced by resuspension, advection, and sedimentation events that contributed organic matter that enhanced bacterial metabolism.     

Comparison of bottom sea-ice algal characteristics from coastal and offshore regions in the Arctic Ocean

Ice-core observations were conducted at three arctic sites in early April 2003: one clean sediment-free site (Chukchi), one sediment-rich site (Beaufort) on landfast sea ice offshore Barrow, and one oceanic Ice Exercise (ICEX) site. Concentrations of inorganic nutrients at coastal sites were similar to but higher than those at the oceanic site. Chlorophyll-a (Chl-a) concentration was much higher in the Chukchi bottom ice than at the other two sites. However, large size Chl-a (>20 μm) dominated (>70%) the bottom-ice algal Chl-a at both the Chukchi and Beaufort sites, whereas the oceanic ICEX site was evenly occupied by large (>20 μm), medium (5–20 μm), and small Chl-a (0.7–5 μm). These in situ data were incorporated into an ocean-ecosystem model. The model results revealed that: (1) strong light attenuation by trapped sediments controlled ice-algal production at the Beaufort site; (2) the peak in ice algae occurred later at the oceanic site than at the Chukchi site because of thicker ice and the consequently reduced amount of light reaching the ice algae at the ice bottom; and (3) maximum production at the oceanic site reached only 10% the level at the Chukchi site because of nutrient limitation.     

Optical fractionation of chlorophyll and primary production for coastal waters of the Southern Ocean

Our objective was to quantify the potential variability in remotely sensed chlorophyll a (Chl a) and primary productivity in coastal waters of the Southern Ocean. From data collected throughout the springs/summers of 1991–1994, we calculated the proportion of water column Chl a and primary productivity within the upper optical attenuation length (K ⁻¹ _par) and the satellite-weighted depth. The temporal variability was resolved every 2–3 days and was observed to be greater within years than between years. Three-year averages (n=223) revealed that 10.2 ± 3.6% of total Chl a and 14.8 ± 6.5% of production occurred within satellite-weighted depth in predominantly Case I waters. The average values were twice as high within K ⁻¹ _par, 24.1 ± 8% of total Chl a and 34 ± 9% of production respectively. Masked in these long-term averages are very large changes occurring on short time scales of seasonal blooms. We observed that the patterns of Chl a vertical distribution within blooms are also subject to taxonomic influence and dependent upon the physiological state of the phytoplankton. Highest proportions of water column Chl a in the first optical depth were measured during the rapid onset of surface cryptophyte blooms each year, i.e. 50% within K ⁻¹ _par and 30% above the satellite-weighted depth. Lowest fractions, 6% and 2% of biomass within K ⁻¹ _par and satellite-weighted depth respectively, were associated with peak bloom conditions independent of taxonomy. Our analyses suggest that satellite-dependent models of Chl a and subsequent chlorophyll-dependent primary production will be challenging to develop for the near-shore Southern Ocean, especially given the potentially high natural variability in the vertical distribution of Chl a driven by physical forcing, the photoadaptive abilities of polar phytoplankton, and taxonomic influences. Accepted: 27 August 1999     

Transmission of parasites in the coastal waters of the Arctic seas and possible effect of climate change

The review deals with the biodiversity, life cycles, distribution and temperature adaptations of parasites circulating in the coastal waters of northern polar seas. Special attention is given to helminths of marine birds, which are the most common parasites in the coastal waters. Among them, the focus is on trematodes. Factors responsible for the impoverished species composition of parasites in the Arctic are analyzed. It is shown that species without free-living larvae in the life cycle have an advantage in this environment. The abundance of cestodes and acanthocephalans in Arctic seabirds is linked with the high proportion of crustaceans in their diet. The phenomenon of nonspecific parasitism (occurrence of parasites in atypical host species) is analyzed from an evolutionary viewpoint. Characteristic features in the spatial distribution of infection of marine coastal invertebrates with parasite larvae are considered, and factors that determine it are specified. The prevalence of infection in intermediate hosts is closely connected with the abundance of final hosts, which makes it possible to estimate the abundance of final hosts in a given region and reveal trends in its changes. Trematodes have a high potential for temperature acclimation. This facilitates their transmission in the northern seas but, on the other hand, makes it unlikely that the transmission process would be intensified upon an increase in summer temperatures resulting from climate warming. However, intensification of transmission may well occur due to the prolongation of the warm season (“transmission window”), which has been predicted and is already observed. It is suggested that warming in the Arctic promotes both the entry of certain “southern” species into the Arctic and the trans-Arctic interpenetration of the North Atlantic and North Pacific parasitic faunas. A case is made for the necessity to broaden the scope of parasitological research in the Arctic and Subarctic, including parasitological monitoring at the reference sites of the sea coast.     

A metaproteomic assessment of winter and summer bacterioplankton from Antarctic Peninsula coastal surface waters

A metaproteomic survey of surface coastal waters near Palmer Station on the Antarctic Peninsula, West Antarctica, was performed, revealing marked differences in the functional capacity of summer and winter communities of bacterioplankton. Proteins from Flavobacteria were more abundant in the summer metaproteome, whereas winter was characterized by proteins from ammonia-oxidizing Marine Group I Crenarchaeota. Proteins prevalent in both seasons were from SAR11 and Rhodobacterales clades of Alphaproteobacteria, as well as many lineages of Gammaproteobacteria. The metaproteome data were used to elucidate the main metabolic and energy generation pathways and transport processes occurring at the microbial level in each season. In summer, autotrophic carbon assimilation appears to be driven by oxygenic photoautotrophy, consistent with high light availability and intensity. In contrast, during the dark polar winter, the metaproteome supported the occurrence of chemolithoautotrophy via the 3-hydroxypropionate/4-hydroxybutyrate cycle and the reverse tricarboxylic acid cycle of ammonia-oxidizing archaea and nitrite-oxidizing bacteria, respectively. Proteins involved in nitrification were also detected in the metaproteome. Taurine appears to be an important source of carbon and nitrogen for heterotrophs (especially SAR11), with transporters and enzymes for taurine uptake and degradation abundant in the metaproteome. Divergent heterotrophic strategies for Alphaproteobacteria and Flavobacteria were indicated by the metaproteome data, with Alphaproteobacteria capturing (by high-affinity transport) and processing labile solutes, and Flavobacteria expressing outer membrane receptors for particle adhesion to facilitate the exploitation of non-labile substrates. TonB-dependent receptors from Gammaproteobacteria and Flavobacteria (particularly in summer) were abundant, indicating that scavenging of substrates was likely an important strategy for these clades of Southern Ocean bacteria. This study provides the first insight into differences in functional processes occurring between summer and winter microbial communities in coastal Antarctic waters, and particularly highlights the important role that ‘dark'' carbon fixation has in winter.     

Phylogenetic Diversity and Ecological Pattern of Ammonia-oxidizing Archaea in the Surface Sediments of the Western Pacific

The phylogenetic diversity of ammonia-oxidizing archaea (AOA) was surveyed in the surface sediments from the northern part of the South China Sea (SCS). The distribution pattern of AOA in the western Pacific was discussed through comparing the SCS with other areas in the western Pacific including Changjiang Estuary and the adjacent East China Sea where high input of anthropogenic nitrogen was evident, the tropical West Pacific Continental Margins close to the Philippines, the deep-sea methane seep sediments in the Okhotsk Sea, the cold deep sea of Northeastern Japan Sea, and the hydrothermal field in the Southern Okinawa Trough. These various environments provide a wide spectrum of physical and chemical conditions for a better understanding of the distribution pattern and diversities of AOA in the western Pacific. Under these different conditions, the distinct community composition between shallow and deep-sea sediments was clearly delineated based on the UniFrac PCoA and Jackknife Environmental Cluster analyses. Phylogenetic analyses showed that a few ammonia-oxidizing archaeal subclades in the marine water column/sediment clade and endemic lineages were indicative phylotypes for some environments. Higher phylogenetic diversity was observed in the Philippines while lower diversity in the hydrothermal vent habitat. Water depth and possibly with other environmental factors could be the main driving forces to shape the phylogenetic diversity of AOA observed, not only in the SCS but also in the whole western Pacific. The multivariate regression tree analysis also supported this observation consistently. Moreover, the functions of current and other climate factors were also discussed in comparison of phylogenetic diversity. The information collectively provides important insights into the ecophysiological requirements of uncultured ammonia-oxidizing archaeal lineages in the western Pacific Ocean.     

Seasonal variation in plankton, submicrometre particles and size-fractionated dissolved organic carbon in Antarctic coastal waters

Concentrations of plankton, suspended particles 0.74–87 μm equivalent spherical diameter and dissolved organic carbon (DOC) were measured from May to February at an Antarctic coastal site. Bacteria-sized particles 0.74–1 μm diameter, and bacterial cells and heterotrophic protists all exhibited a seasonal minimum during winter and maxima in summer. Bacteria composed <10% of the bacteria-sized particles. Release of autotrophic protists from the ice caused water column biomass of autotrophs to reach maximum concentrations in October and November, but maximum cell concentration in the water column was reached in January. Microheterotroph biomass weakly reflected the release of the ice algal community but reached maximum concentration during the water column bloom in January. Total DOC concentrations varied from 0.36 mg C l⁻¹ in July to 3.10 mg C l⁻¹ in October, with a yearly average of 1.51 mg C l⁻¹. Ultrafiltration of DOC revealed that the molecular weight composition of the DOC differed greatly through the year. DOC <5 kDa molecular weight reached a maximum of 1.25 mg C l⁻¹ in October and accounted for up to 60% of total DOC in July. Concentrations of high molecular weight DOC (>100 kDa) were highest in July and November, with the DOC (100 kDa–0.5 μm) fraction reaching a maximum of 1.22 mg C l⁻¹ in November and composing 82% of the total DOC in January. Wet chemical oxidation and high-temperature catalytic oxidation organic carbon analyses were compared. Good correlation was observed between methods during summer but no significant correlation existed in winter, indicating that winter DOC may be refractory. Accepted: 21 March 2000     

Bacteriophages in Arctic and Antarctic low-temperature systems

Comparative analysis of the presence of bacteriophages was carried out for the water column of a permanently ice-covered, extremely oligotrophic Lake Untersee (East Antarctica) and the ancient ice wedge of the Mamontova Gora outcrop (Aldan River, Central Yakutia). Microscopy revealed bacteriophages in the Mamontova Gora ice samples and in the lysates of the pure cultures of phage-sensitive bacteria isolated from the same samples. Bacteriophages isolated from these cultures were filamentous and interacted with bacteria as moderate (lysogenic) phages. A similar filamentous bacteriophage was isolated from the Lake Untersee water column. The highest morphological diversity of bacteriophages was revealed by microscopy in the oxic Lake Untersee water column in the chemocline zone (70–76 m) and in the sulfide layer (85 m). Detection of similar filamentous bacteriophages in a relic ice sample and in the samples from Antarctic Lake Untersee indicate wide occurrence of bacteriophages and lysogeny in microbial communities of low-temperature ecosystems.     

Hydroid epifaunal communities in Arctic coastal waters (Svalbard): effects of substrate characteristics

The knowledge of cryptic epifaunal groups in the Arctic is far from complete mostly due to logistic difficulties. Only recently, advances in sample collection using SCUBA diving techniques have enabled to explore delicate hydroid fauna from shallow waters. This study is the first attempt to examine the relationship between substrate property (such as size of rock, morphological characteristics of algal or bryozoan host) and hydroid community composition and diversity in the Arctic. Samples of substrates for hydroid attachment including rocks, algae, bryozoans and other hydrozoans were collected around the Svalbard. Examination revealed no substrate-specific species. The substrate property did not have a strong influence on hydroid community. Both species composition and richness were not related to colonized rock surface area and to morphological characteristic of algal host. Therefore, results indicate the opportunistic nature of hydroid fauna in terms of substrate preference. However, the presence or absence of hydroids depended on the surface area of rocky substrate. Hydroids were more often present on rocks of larger surface area. Erect hydroids and bryozoans were important attachment surface for stolonal hydroids.     

Lipids and fatty acids in Clione limacina and Limacina helicina in Svalbard waters and the Arctic Ocean: trophic implications

Lipid class and fatty acid compositions were determined in Limacina helicina and Clione limacina from an Arctic fjord and the marginal ice zone around Svalbard. C. limacina had higher levels of neutral lipids, including both alkyldiacylglycerols (ADG) and triacylglycerols (TAG), than L. helicina, which contained mainly TAG. However, considerable heterogeneity in the lipid classes and their fatty acids/alcohols were observed in C. limacina in that only two out of the seven specimens analysed were lipid-rich and contained both ADG and TAG, the others having only low percentages of TAG. In specimens of C. limacina containing ADG, 15:0 and 17:1n-8 were prominent fatty acids in both ADG and TAG. The fatty acids of the TAG of L. helicina were variable but 15:0 and 17:1n-8 were absent. We consider the heterogeneity in the fatty acid compositions of L. helicina to reflect temporal and spatial variability in the animals' predominantly phytoplanktonic and particulate diet, which occasionally includes small copepods. We further consider L. helicina to be the prime food for C. limacina and the noticeable amounts of 22:1 found in one sample of C. limacina to reflect significant input of Calanus either directly or indirectly through their prime food, L. helicina. We view the heterogeneity in the fatty acid compositions of both L. helicina and C. limacina, as well as the ability of C. limacina to biosynthesise WE, ADG, 15:0, and 17:1n-8, as adaptations to a large variation of food availability that enables C. limacina to synthesise lipids rapidly and flexibly. Thus, the lipid biochemistry of C. limacina is important in enabling the species to thrive in strong pulses in polar systems. Revised: 16 August 2000 / Accepted: 17 August 2000     

Aerobic microorganisms associated with free-ranging bottlenose dolphins in coastal Gulf of Mexico and Atlantic Ocean waters

Our abilities to assess health risks to free-ranging dolphin populations, to treat live-stranded or captive dolphins, and to evaluate the risks of disease transmission between humans and dolphins have suffered from a lack of basic information on microorganisms associated with normal, presumably healthy free-ranging individuals. In order to provide these data, we sampled free-ranging bottlenose dolphins (Tursiops truncatus) off Florida, Texas, and North Carolina during 1990-2002. Blowhole and anal/fecal samples yielded 1,871 bacteria and yeast isolates and included 85 different species or groups of organisms. Vibrios, unidentified pseudomonads, Escherichia coli, Staphylococcus spp., and a large group of nonfermenting gram-negative bacteria represented >50% of isolates. Vibrio alginolyticus and Vibrio damsela were the most commonly recovered bacteria from both anal/fecal and blowhole samples. Many organisms occurred sporadically in dolphins that were sampled repeatedly, but some were consistently isolated from individual animals and may indicate the carrier state. Vibrios were common, but some geographic variability in the presence of these and other organisms was noted. Potential pathogens of significance to humans and other animals were recovered.     

Feeding and diel vertical migration cycles of Metridia gerlachei (Giesbrecht) in coastal waters of the Antarctic Peninsula

Diel vertical migration and feeding cycles of adult female Metridia gerlachei in the upper 290 m of a 335-m water column were measured during a total of 65 h in two periods of early summer (Dec 20–21 and Dec 25–26, 1991). Samples collected in eight depth strata by 35 MOCNESS tows (333-m mesh) were analyzed for abundance and mean individual gut pigment content. Most of the copepod population was concentrated in a 50-m depth interval at all times. Feeding began simultaneously with nocturnal ascent from a depth of 200–250 m at 18:00 h (local time), when the relative change in ambient light intensity was greatest. Ingestion rate increased exponentially (k_i = 0.988 h^–1) at double the gut evacuation rate (k_e = 0.488 h^–1) as the population moved upward at 22.3–26.5 m h^–1 through increasing concentrations of particulate chlorophyll-a. Although the bulk of the population did not move to depths shallower than 50 m, and began its downward migration at a rate of 20.8–31.7 mh^–1 in complete darkness, individual females continued to make brief excursions into chlorophyll-rich surface waters (4–8 g l^–1) during the first few hours of population descent. Ingestion rate diminished abruptly by one order of magnitude (k_i = 0.068 h^–1) at dawn ( 0330 h). Within four more hours, the population had reached its daytime depth and gut pigment content remained constant at a minimum value until the next migration cycle. No feeding appeared to take place at depth during the day. Ingestion by M. gerlachei females removed < 4% of daily primary production, with only 20% of this amount being removed from surface waters by active vertical transport.