Role of macroscopic particles in deep-sea oxygen consumption.
ABSTRACT: Macroscopic particles (>500 mum), including marine snow, large migrating zooplankton, and their fast-sinking fecal pellets, represent primary vehicles of organic carbon flux from the surface to the deep sea. In contrast, freely suspended microscopic particles such as bacteria and protists do not sink, and they contribute the largest portion of metabolism in the upper ocean. In bathy- and abyssopelagic layers of the ocean (2,000-6,000 m), however, microscopic particles may not dominate oxygen consumption. In a section across the tropical Atlantic, we show that macroscopic particle peaks occurred frequently in the deep sea, whereas microscopic particles were barely detectable. In 10 of 17 deep-sea profiles (>2,000 m depth), macroscopic particle abundances were more strongly cross-correlated with oxygen deficits than microscopic particles, suggesting that biomass bound to large particles dominates overall deep-sea metabolism.
Project description:Particles are the major vector for the transfer of carbon from the upper ocean to the deep sea. However, little is known about their abundance, composition and role at depths greater than 2000 m. We present the first number-size spectrum of bathy- and abyssopelagic particles to a depth of 5500 m based on surveys performed with a custom-made holographic microscope. The particle spectrum was unusual in that particles of several millimetres in length were almost 100 times more abundant than expected from the number spectrum of smaller particles, thereby meeting the definition of "dragon kings." Marine snow particles overwhelmingly contributed to the total particle volume (95-98%). Approximately 1/3 of the particles in the dragon-king size domain contained large amounts of transparent exopolymers with little ballast, which likely either make them neutrally buoyant or cause them to sink slowly. Dragon-king particles thus provide large volumes of unique microenvironments that may help to explain discrepancies in deep-sea biogeochemical budgets.
Project description:Particle-attached (PA) and free-living (FL) microorganisms play significant but different roles in mineralization of organic matter (OM) in the ocean. Currently, little is known about PA and FL microbial communities in bathyal and abyssal pelagic waters, and understanding of their diversity and distribution in the water column and their interactions with environmental factors in the trench area is limited. We investigated for the first time the variations of abundance and diversities of the PA and FL bacterial communities in the epi-, bathy-, and abyssopelagic zones of the New Britain Trench (NBT). The PA communities showed decreasing species richness but increasing relative abundance with depth, suggesting the increasing ecological significance of the PA bacteria in the deep ocean. The abundance and diversity of PA and FL bacterial communities in the NBT water column appeared to be shaped by different sets of environment factors, which might be related to different micro-niches of the two communities. Analysis on species distribution suggested that the differences between PA and FL bacteria communities mainly resulted from the different relative abundance of the "shared taxa" in the two types of communities. These findings provide valuable information for understanding the relative ecological roles of the PA and FL bacterial communities and their interactions with environmental factors in different pelagic zones along the vertical profile of the NBT water column.
Project description:The sinking of organic particles formed in the photic layer is a main vector of carbon export into the deep ocean. Although sinking particles are heavily colonized by microbes, so far it has not been explored whether this process plays a role in transferring prokaryotic diversity from surface to deep oceanic layers. Using Illumina sequencing of the 16S rRNA gene, we explore here the vertical connectivity of the ocean microbiome by characterizing marine prokaryotic communities associated with five different size fractions and examining their compositional variability from surface down to 4,000 m across eight stations sampled in the Atlantic, Pacific, and Indian Oceans during the Malaspina 2010 Expedition. Our results show that the most abundant prokaryotes in the deep ocean are also present in surface waters. This vertical community connectivity seems to occur predominantly through the largest particles because communities in the largest size fractions showed the highest taxonomic similarity throughout the water column, whereas free-living communities were more isolated vertically. Our results further suggest that particle colonization processes occurring in surface waters determine to some extent the composition and biogeography of bathypelagic communities. Overall, we postulate that sinking particles function as vectors that inoculate viable particle-attached surface microbes into the deep-sea realm, determining to a considerable extent the structure, functioning, and biogeography of deep ocean communities.
Project description:Greater diversity of eukaryotic phytoplankton than expected has been revealed recently through molecular techniques, but little is known about their temporal dynamics or fate in the open ocean. Here, we examined size-fractionated eukaryotic phytoplankton communities from the surface to abyssopelagic zone (5,000 m) throughout the year, by tracking sequence variants of the 18S rRNA gene in the western subtropical North Pacific. The oceanographic conditions were divided into two periods, stratification and mixing, between which the surface phytoplankton community differed. During the mixing period, the abundance of large phytoplankton (?3 ?m) increased, with diatoms and putative Pseudoscourfieldia marina dominating this fraction. Picophytoplankton (<3 ?m) also increased during the mixing period and were dominated by Mamiellophyceae. Taxa belonging to prasinophytes (including Ps. marina and Mamiellophyceae) were observed in the epipelagic zone throughout the year, and thus likely seeded the seasonal bloom that occurred during the mixing period. In contrast, diatoms observed during the mixing period mostly represented taxa unique to that period, including coastal species. Numerical particle backtracking experiments indicated that water masses in the surface layer could be transported from coastal areas to the study site. Gene sequences of coastal diatoms were present in the abyssopelagic zone. Therefore, allochthonous species drove the seasonal bloom and could be transported to deep waters. In the abyssopelagic zone, the relative abundance of Ps. marina in deep waters was similar to or higher than that of diatoms during the mixing period. Among picophytoplankton, Mamiellophyceae made up a significant fraction in the abyssopelagic zone, suggesting that prasinophytes are also involved in carbon export. Our molecular survey showed that these previously overlooked phytoplankton species could contribute significantly to the seasonal bloom and biological pump in the subtropical open ocean.
Project description:Picoeukaryotes play prominent roles in the biogeochemical cycles in marine ecosystems. However, their molecular diversity studies have been confined in marine surface waters or shallow coastal sediments. Here, we investigated the diversity and metabolic activity of picoeukaryotic communities at depths ranging from the surface to the abyssopelagic zone in the western Pacific Ocean above the north and south slopes of the Mariana Trench. This was achieved by amplifying and sequencing the V4 region of both 18S ribosomal DNA and cDNA using Illumina HiSeq sequencing. Our study revealed: (1) Four super-groups (i.e., Alveolata, Opisthokonta, Rhizaria and Stramenopiles) dominated the picoeukaryote assemblages through the water column, although they accounted for different proportions at DNA and cDNA levels. Our data expand the deep-sea assemblages from current bathypelagic to abyssopelagic zones. (2) Using the cDNA-DNA ratio as a proxy of relative metabolic activity, the highest activity for most subgroups was usually found in the mesopelagic zone; and (3) Population shift along the vertical scale was more prominent than that on the horizontal differences, which might be explained by the sharp physicochemical gradients along the water depths. Overall, our study provides a better understanding of the diversity and metabolic activity of picoeukaryotes in water columns of the deep ocean in response to varying environmental conditions.
Project description:Sinking particles are a critical conduit for the export of organic material from surface waters to the deep ocean. Despite their importance in oceanic carbon cycling and export, little is known about the biotic composition, origins, and variability of sinking particles reaching abyssal depths. Here, we analyzed particle-associated nucleic acids captured and preserved in sediment traps at 4,000-m depth in the North Pacific Subtropical Gyre. Over the 9-month time-series, Bacteria dominated both the rRNA-gene and rRNA pools, followed by eukaryotes (protists and animals) and trace amounts of Archaea. Deep-sea piezophile-like Gammaproteobacteria, along with Epsilonproteobacteria, comprised >80% of the bacterial inventory. Protists (mostly Rhizaria, Syndinales, and ciliates) and metazoa (predominantly pelagic mollusks and cnidarians) were the most common sinking particle-associated eukaryotes. Some near-surface water-derived eukaryotes, especially Foraminifera, Radiolaria, and pteropods, varied greatly in their abundance patterns, presumably due to sporadic export events. The dominance of piezophile-like Gammaproteobacteria and Epsilonproteobacteria, along with the prevalence of their nitrogen cycling-associated gene transcripts, suggested a central role for these bacteria in the mineralization and biogeochemical transformation of sinking particulate organic matter in the deep ocean. Our data also reflected several different modes of particle export dynamics, including summer export, more stochastic inputs from the upper water column by protists and pteropods, and contributions from sinking mid- and deep-water organisms. In total, our observations revealed the variable and heterogeneous biological origins and microbial activities of sinking particles that connect their downward transport, transformation, and degradation to deep-sea biogeochemical processes.
Project description:The Japan Trench is located under the eutrophic Northwestern Pacific while the Mariana Trench that harbors the unique hadal planktonic biosphere is located under the oligotrophic Pacific. Water samples from the sea surface to just above the seafloor at a total of 11 stations including a trench axis station, were investigated several months after the Tohoku Earthquake in March 2011. High turbidity zones in deep waters were observed at most of the sampling stations. The small subunit (SSU) rRNA gene community structures in the hadal waters (water depths below 6000 m) at the trench axis station were distinct from those in the overlying meso-, bathy and abyssopelagic waters (water depths between 200 and 1000 m, 1000 and 4000 m, and 4000 and 6000 m, respectively), although the SSU rRNA gene sequences suggested that potential heterotrophic bacteria dominated in all of the waters. Potential niche separation of nitrifiers, including ammonia-oxidizing archaea (AOA), was revealed by quantitative PCR analyses. It seems likely that Nitrosopumilus-like AOAs respond to a high flux of electron donors and dominate in several zones of water columns including shallow and very deep waters. This study highlights the effects of suspended organic matter, as induced by seafloor deformation, on microbial communities in deep waters and confirm the occurrence of the distinctive hadal biosphere in global trench environments hypothesized in the previous study.
Project description:The deep sea is one of the largest but least understood ecosystems on earth. Knowledge about the diversity and distribution patterns as well as drivers of microbial eukaryote (including ciliates) along the water column, particularly below the photic zone, is scarce. In this study, we investigated the diversity of pelagic ciliates, the main group of marine microeukaryotes, their vertical distribution from the surface to the abyssopelagic zone, as well as their horizontal distribution over a distance of 1,300 km in the Western Pacific Ocean, using high-throughput DNA and cDNA (complementary DNA) sequencing. No distance-decay relationship could be detected along the horizontal scale; instead, a distinct vertical distribution within the ciliate communities was revealed. The alpha diversity of the ciliate communities in the deep chlorophyll maximum (DCM) and the 200 m layer turned out to be significantly higher compared with the other water layers. The ciliate communities in the 200 m water layer appeared to be more similar to those in deeper layers from 1,000 m to about 5,000 m than to the surface and DCM ciliate communities. Dominant species in the bathypelagic and abyssopelagic zone, particularly some parasites, were also detected in the 200 m layer, but were almost absent in the surface layer. The 200 m layer, therefore, seems to be an important "species bank" for deep ocean layers. Statistical analyses further revealed significant effects of temperature and chlorophyll a on the partitioning of ciliate diversity, indicating that environmental factors are a stronger force in shaping marine pelagic ciliate communities than the geographic distance.
Project description:Despite numerous studies on marine prokaryotes, the vertical distribution patterns of bacterial community, either on the taxonomic composition or the functional structure, remains relatively unexplored. Using HiSeq-derived 16S rRNA data, the depth-related distribution patterns of taxonomic diversity and functional structure predicted from diversity data in the water column and sediments of the Western Pacific Ocean were explored. The OTU richness declined along the water column after peaking between 100 to 200 m deep. Relative abundance of Cyanobacteria and SAR11 decreased significantly with depth, while Actinobacteria and Gammaproteobacteria increased. This clearly mirrors the vertical distribution pattern of the predicted functional composition with the shift between phototrophic to chemoheterotrophic groups from the surface to the deeper layers. In terms of community composition and functional structure, the epipelagic zone differed from other deeper ones (i.e., meso-, bathy-, and abyssopelagic zones) where no obvious differences were detected. For the epipelagic zone, temperature, dissolved oxygen, and salinity were recognized as the crucial factors shaping both community composition and the functional structure of bacteria. Compared with water samples, benthic sediment samples harbored unexpectedly higher read abundance of Proteobacteria, presenting distinguishable taxonomic and functional compositions. This study provides novel knowledge on the vertical distribution of bacterial taxonomic and functional compositions in the western Pacific.
Project description:Viral abundance in deep-sea environments is high. However, the biological, ecological and biogeochemical roles of viruses in the deep sea are under debate. In the present study, microcosm incubations of deep-sea bacterioplankton (2,000?m deep) with normal and reduced pressure of viral lysis were conducted in the western Pacific Ocean. We observed a negative effect of viruses on prokaryotic abundance, indicating the top-down control of bacterioplankton by virioplankton in the deep-sea. The decreased bacterial diversity and a different bacterial community structure with diluted viruses indicate that viruses are sustaining a diverse microbial community in deep-sea environments. Network analysis showed that relieving viral pressure decreased the complexity and clustering coefficients but increased the proportion of positive correlations for the potentially active bacterial community, which suggests that viruses impact deep-sea bacterioplankton interactions. Our study provides experimental evidences of the crucial role of viruses in microbial ecology and biogeochemistry in deep-sea ecosystems.