LIMNOLOGY AND OCEANOGRAPHY

Bacterial biogeography of the Indian Ocean
Brock ML, Larkin AA, Raes EJ and Martiny AC
Historically, our understanding of bacterial ecology in the Indian Ocean has been limited to regional studies that place emphasis on community structure and function within oxygen minimum zones. Thus, bacterial community dynamics across the wider Indian Ocean are largely undescribed. As part of Bio-GO-SHIP, we sequenced the 16S rRNA gene from 465 samples collected on sections I07N and I09N. We found that (i) there were 23 distinct bioregions within the Indian Ocean, (ii) the southeastern gyre had the largest gradient in bacterial alpha-diversity, (iii) the Indian Ocean surface microbiome was primarily composed of a core set of taxa, and (iv) bioregions were characterized by transitions in physical and geochemical conditions. Overall, we showed that bacterial community structure spatially delineated the surface Indian Ocean and that these microbially-defined regions were reflective of subtle ocean physical and geochemical gradients. Therefore, incorporating metrics of in-situ microbial communities into marine ecological regions traditionally defined by remote sensing will improve our ability to delineate warm, oligotrophic regions.
Pulses of labile carbon cause transient decoupling of fermentation and respiration in permeable sediments
Nauer PA, Kessler AJ, Hall P, Popa ME, Ten Hietbrink S, Hutchinson T, Wong WW, Attard K, Glud RN, Greening C and Cook PLM
Dihydrogen (H) is an important intermediate in anaerobic microbial processes, and concentrations are tightly controlled by thermodynamic limits of consumption and production. However, recent studies reported unusual H accumulation in permeable marine sediments under anoxic conditions, suggesting decoupling of fermentation and sulfate reduction, the dominant respiratory process in anoxic permeable marine sediments. Yet, the extent, prevalence and potential triggers for such H accumulation and decoupling remain unknown. We surveyed H concentrations in situ at different settings of permeable sand and found that H accumulation was only observed during a coral spawning event on the Great Barrier Reef. A flume experiment with organic matter addition to the water column showed a rapid accumulation of hydrogen within the sediment. Laboratory experiments were used to explore the effect of oxygen exposure, physical disturbance and organic matter inputs on H accumulation. Oxygen exposure had little effect on H accumulation in permeable sediments suggesting both fermenters and sulfate reducers survive and rapidly resume activity after exposure to oxygen. Mild physical disturbance mimicking sediment resuspension had little effect on H accumulation; however, vigorous shaking led to a transient accumulation of H and release of dissolved organic carbon suggesting mechanical disturbance and cell destruction led to organic matter release and transient decoupling of fermenters and sulfate reducers. In summary, the highly dynamic nature of permeable sediments and its microbial community allows for rapid but transient decoupling of fermentation and respiration after a C pulse, leading to high H levels in the sediment.
Aerobic methane synthesis and dynamics in a river water environment
Alowaifeer AM, Wang Q, Bothner B, Sibert RJ, Joye SB and McDermott TR
Reports of aerobic biogenic methane () have generated new views about sources in nature. We examine this phenomenon in the free-flowing Yellowstone river wherein concentrations were tracked as a function of environmental conditions, phototrophic microorganisms (using chlorophyll , Chl , as proxy), as well as targeted methylated amines known to be associated with this process. was positively correlated with temperature and Chl , although diurnal measurements showed concentrations were greatest during the night and lowest during maximal solar irradiation. efflux from the river surface was greater in quiescent edge waters (71-94 mol m d) than from open flowing current (~ 57 mol m d). Attempts to increase flux by disturbing the benthic environment in the quiescent water directly below (~ 1.0 m deep) or at varying distances (0-5 m) upstream of the flux chamber failed to increase surface flux. Glycine betaine (GB), dimethylamine and methylamine (MMA) were observed throughout the summer-long study, increasing during a period coinciding with a marked decline in Chl , suggesting a lytic event led to their release; however, this did not correspond to increased concentrations. Spiking river water with GB or MMA yielded significantly greater than nonspiked controls, illustrating the metabolic potential of the river microbiome. In summary, this study provides evidence that: (1) phototrophic microorganisms are involved in synthesis in a river environment; (2) the river microbiome possesses the metabolic potential to convert methylated amines to ; and (3) river concentrations are dynamic diurnally as well as during the summer active months.
Diazotrophy modulates cyanobacteria stoichiometry through functional traits that determine bloom magnitude and toxin production
Wagner ND, Osburn FS, Taylor RB, Back JA, Chambliss CK, Brooks BW and Scott JT
Harmful cyanobacterial blooms are an increasing threat to water quality. The interactions between two eco-physiological functional traits of cyanobacteria, diazotrophy (nitrogen (N)-fixation) and N-rich cyanotoxin synthesis, have never been examined in a stoichiometric explicit manner. We explored how a gradient of resource N:phosphorus (P) affects the biomass, N, P stoichiometry, light-harvesting pigments, and cylindrospermopsin production in a N-fixing cyanobacterium, . Low N:P cultures produced the same biomass as populations grown in high N:P cultures. The biomass accumulation determined by carbon, indicated low N:P cultures did not have a N-fixation growth tradeoff, in contrast to some other diazotrophs that maintain stoichiometric N homeostasis at the expense of growth. However, N-fixing populations produced less particulate cylindrospermopsin and had undetectable dissolved cylindrospermopsin compared to non-N-fixing populations. The pattern of low to high cyanotoxin cell quotas across an N:P gradient in the diazotrophic cylindrospermopsin producer is similar to the cyanotoxin cell quota response in non-diazotrophic cyanobacteria. We suggest that diazotrophic cyanobacteria may be characterized into two broad functional groups, the N-storage-strategists and the growth-strategists, which use N-fixation differently and may determine patterns of bloom magnitude and toxin production in nature.
Carbon content, carbon fixation yield and dissolved organic carbon release from diverse marine nitrifiers
Bayer B, McBeain K, Carlson CA and Santoro AE
Nitrifying microorganisms, including ammonia-oxidizing archaea, ammonia-oxidizing bacteria, and nitrite-oxidizing bacteria, are the most abundant chemoautotrophs in the ocean and play an important role in the global carbon cycle by fixing dissolved inorganic carbon (DIC) into biomass. The release of organic compounds by these microbes is not well quantified, but may represent an as-yet unaccounted source of dissolved organic carbon (DOC) available to marine food webs. Here, we provide measurements of cellular carbon and nitrogen quotas, DIC fixation yields and DOC release of 10 phylogenetically diverse marine nitrifiers. All investigated strains released DOC during growth, representing on average 5-15% of the fixed DIC. Changes in substrate concentration and temperature did not affect the proportion of fixed DIC released as DOC, but release rates varied between closely related species. Our results also indicate previous studies may have underestimated DIC fixation yields of marine nitrite oxidizers due to partial decoupling of nitrite oxidation from CO fixation, and due to lower observed yields in artificial compared to natural seawater medium. The results of this study provide critical values for biogeochemical models of the global carbon cycle, and help to further constrain the implications of nitrification-fueled chemoautotrophy for marine food-web functioning and the biological sequestration of carbon in the ocean.
Planktonic microbial communities from microbialite-bearing lakes sampled along a salinity-alkalinity gradient
Iniesto M, Moreira D, Benzerara K, Reboul G, Bertolino P, Tavera R and López-García P
Continental freshwater systems are particularly vulnerable to environmental variation. Climate change-induced desertification and the anthropogenic exploitation of hydric resources result in the progressive evaporation and salinization of inland water bodies in many areas of the globe. However, how this process impacts microbial communities and their activities in biogeochemical cycles is poorly known. Here, we take a space-for-time substitution approach and characterize the prokaryotic and eukaryotic microbial communities of two planktonic cell-size fractions (0.2-5 m and 5-30 m) from lakes of diverse trophic levels sampled along a salinity-alkalinity gradient located in the Trans-Mexican Volcanic Belt (TMVB). We applied a 16S/18S rRNA gene metabarcoding strategy to determine the microbial community composition of 54 samples from 12 different lakes, from the low-salinity lake Zirahuén to the hypersaline residual ponds of Rincón de Parangueo. Except for systems at both extremes of the salinity gradient, most lakes along the evaporation trend bear actively forming microbialites, which harbor microbial communities clearly distinct from those of plankton. Several lakes were sampled in winter and late spring and the crater lakes Alchichica and Atexcac were sampled across the water column. Physicochemical parameters related to salinity-alkalinity were the most influential drivers of microbial community structure whereas trophic status, depth, or season were less important. Our results suggest that climate change and anthropogenic-induced hydric deficit could significantly affect microbial communities, potentially altering ecosystem functioning.
Impact of local rivers on coastal acidification
Savoie AM, Moody A, Gilbert M, Dillon KS, Howden SD, Shiller AM and Hayes CT
Coastal ecosystems are highly dynamic areas for carbon cycling and are likely to be negatively impacted by increasing ocean acidification. This research focused on dissolved inorganic carbon (DIC) and total alkalinity (TA) in the Mississippi Sound to understand the influence of local rivers on coastal acidification. This area receives large fluxes of freshwater from local rivers, in addition to episodic inputs from the Mississippi River through a human-built diversion, the Bonnet Carré Spillway. Sites in the Sound were sampled monthly from August 2018 to November 2019 and weekly from June to August 2019 in response to an extended spillway opening. Prior to the 2019 spillway opening, the contribution of the local, lower alkalinity rivers to the Sound may have left the study area more susceptible to coastal acidification during winter months, with aragonite saturation states (Ω) < 2. After the spillway opened, despite a large increase in TA throughout the Sound, aragonite saturation states remained low, likely due to hypoxia and increased CO concentrations in subsurface waters. Increased Mississippi River input could represent a new normal in the Sound's hydrography during spring and summer months. The spillway has been utilized more frequently over the last two decades due to increasing precipitation in the Mississippi River watershed, which is primarily associated with climate change. Future increases in freshwater discharge and the associated declines in salinity, dissolved oxygen, and Ω in the Sound will likely be detrimental to oyster stocks and the resilience of similar ecosystems to coastal acidification.
Emerging harmful algal blooms caused by distinct seasonal assemblages of a toxic diatom
Sterling AR, Kirk RD, Bertin MJ, Rynearson TA, Borkman DG, Caponi MC, Carney J, Hubbard KA, King MA, Maranda L, McDermith EJ, Santos NR, Strock JP, Tully EM, Vaverka SB, Wilson PD and Jenkins BD
Diatoms in the genus produce the neurotoxin domoic acid. Domoic acid bioaccumulates in shellfish, causing illness in humans and marine animals upon ingestion. In 2017, high domoic acid levels in shellfish meat closed shellfish harvest in Narragansett Bay, Rhode Island for the first and only time in history, although abundant have been observed for over 60 years To investigate whether an environmental factor altered endemic physiology or new domoic acid-producing strain(s) were introduced to Narragansett Bay, we conducted weekly sampling from 2017 to 2019 and compared closure samples. Plankton-associated domoic acid was quantified by LC-MS/MS and spp. were identified using a taxonomically improved high-throughput rDNA sequencing approach. Comparison with environmental data revealed a detailed understanding of domoic acid dynamics and seasonal multi-species assemblages. Plankton-associated domoic acid was low throughout 2017-2019, but recurred in fall and early summer maxima. Fall domoic acid maxima contained known toxic species as well as a novel genotype. Summer domoic acid maxima included fewer species but also known toxin producers Most 2017 closure samples contained the particularly concerning toxic species, , which also appeared infrequently during 2017-2019. Recurring assemblages were driven by seasonal temperature changes, and plankton-associated domoic acid correlated with low dissolved inorganic nitrogen. Thus, the Narragansett Bay closures were likely caused by both resident assemblages that become toxic depending on nutrient status as well as the episodic introductions of toxic species from oceanographic and climatic shifts.
Machine learning techniques to characterize functional traits of plankton from image data
Orenstein EC, Ayata SD, Maps F, Becker ÉC, Benedetti F, Biard T, de Garidel-Thoron T, Ellen JS, Ferrario F, Giering SLC, Guy-Haim T, Hoebeke L, Iversen MH, Kiørboe T, Lalonde JF, Lana A, Laviale M, Lombard F, Lorimer T, Martini S, Meyer A, Möller KO, Niehoff B, Ohman MD, Pradalier C, Romagnan JB, Schröder SM, Sonnet V, Sosik HM, Stemmann LS, Stock M, Terbiyik-Kurt T, Valcárcel-Pérez N, Vilgrain L, Wacquet G, Waite AM and Irisson JO
Plankton imaging systems supported by automated classification and analysis have improved ecologists' ability to observe aquatic ecosystems. Today, we are on the cusp of reliably tracking plankton populations with a suite of lab-based and in situ tools, collecting imaging data at unprecedentedly fine spatial and temporal scales. But these data have potential well beyond examining the abundances of different taxa; the individual images themselves contain a wealth of information on functional traits. Here, we outline traits that could be measured from image data, suggest machine learning and computer vision approaches to extract functional trait information from the images, and discuss promising avenues for novel studies. The approaches we discuss are data agnostic and are broadly applicable to imagery of other aquatic or terrestrial organisms.
Length, width, shape regularity, and chain structure: time series analysis of phytoplankton morphology from imagery
Sonnet V, Guidi L, Mouw CB, Puggioni G and Ayata SD
Functional traits are increasingly used to assess changes in phytoplankton community structure and to link individual characteristics to ecosystem functioning. However, they are usually inferred from taxonomic identification or manually measured for each organism, both time consuming approaches. Instead, we focus on high throughput imaging to describe the main temporal variations of morphological changes of phytoplankton in Narragansett Bay, a coastal time-series station. We analyzed a 2-yr dataset of morphological features automatically extracted from continuous imaging of individual phytoplankton images (~ 105 million images collected by an Imaging FlowCytobot). We identified synthetic morphological traits using multivariate analysis and revealed that morphological variations were mainly due to changes in length, width, shape regularity, and chain structure. Morphological changes were especially important in winter with successive peaks of larger cells with increasing complexity and chains more clearly connected. Small nanophytoplankton were present year-round and constituted the base of the community, especially apparent during the transitions between diatom blooms. High inter-annual variability was also observed. On a weekly timescale, increases in light were associated with more clearly connected chains while more complex shapes occurred at lower nitrogen concentrations. On an hourly timescale, temperature was the determinant variable constraining cell morphology, with a general negative influence on length and a positive one on width, shape regularity, and chain structure. These first insights into the phytoplankton morphology of Narragansett Bay highlight the possible morphological traits driving the phytoplankton succession in response to light, temperature, and nutrient changes.
Quantification of microcystin production and biodegradation rates in the western basin of Lake Erie
Chaffin JD, Westrick JA, Furr E, Birbeck JA, Reitz LA, Stanislawczyk K, Li W, Weber PK, Bridgeman TB, Davis TW and Mayali X
Cyanobacterial biomass forecasts currently cannot predict the concentrations of microcystin, one of the most ubiquitous cyanotoxins that threaten human and wildlife health globally. Mechanistic insights into how microcystin production and biodegradation by heterotrophic bacteria change spatially and throughout the bloom season can aid in toxin concentration forecasts. We quantified microcystin production and biodegradation during two growth seasons in two western Lake Erie sites with different physicochemical properties commonly plagued by summer blooms. Microcystin production rates were greater with elevated nutrients than under ambient conditions and were highest nearshore during the initial phases of the bloom, and production rates were lower in later bloom phases. We examined biodegradation rates of the most common and toxic microcystin by adding extracellular stable isotope-labeled microcystin-LR (1 g L), which remained stable in the abiotic treatment (without bacteria) with minimal adsorption onto sediment, but strongly decreased in all unaltered biotic treatments, suggesting biodegradation. Greatest biodegradation rates (highest of -8.76 d, equivalent to the removal of 99.98% in 18 h) were observed during peak bloom conditions, while lower rates were observed with lower cyanobacteria biomass. Cell-specific nitrogen incorporation from microcystin-LR by nanoscale imaging mass spectrometry showed that a small percentage of the heterotrophic bacterial community actively degraded microcystin-LR. Microcystin production and biodegradation rates, combined with the microcystin incorporation by single cells, suggest that microcystin predictive models could be improved by incorporating toxin production and biodegradation rates, which are influenced by cyanobacterial bloom stage (early vs. late bloom), nutrient availability, and bacterial community composition.
Aerobic and anaerobic methane oxidation in a seasonally anoxic basin
Steinsdóttir HGR, Schauberger C, Mhatre S, Thamdrup B and Bristow LA
Shallow coastal waters are dynamic environments that dominate global marine methane emissions. Particularly high methane concentrations are found in seasonally anoxic waters, which are spreading in eutrophic coastal systems, potentially leading to increased methane emissions to the atmosphere. Here we explore how the seasonal development of anoxia influenced methane concentrations, rates of methane oxidation, and the community composition of methanotrophs in the shallow eutrophic water column of Mariager Fjord, Denmark. Our results show the development of steep concentration gradients toward the oxic-anoxic interface as methane accumulated to 1.4 M in anoxic bottom waters. Yet, the fjord possessed an efficient microbial methane filter near the oxic-anoxic interface that responded to the increasing methane flux. In experimental incubations, methane oxidation near the oxic-anoxic interface proceeded both aerobically and anaerobically with nearly equal efficiency reaching turnover rates as high as 0.6 and 0.8 d, respectively, and was seemingly mediated by members of the Methylococcales belonging to the Deep Sea-1 clade. Throughout the period, both aerobic and anaerobic methane oxidation rates were high enough to consume the estimated methane flux. Thus, our results indicate that seasonal anoxia did not increase methane emissions.
The influence of particle concentration and bulk characteristics on polarized oceanographic lidar measurements
Collister BL, Zimmerman RC, Sukenik CI, Balch WM and Hill VJ
Oceanographic lidar measurements of the linear depolarization ratio, , contain information on the bulk characteristics of marine particles that could improve our ability to study ocean biogeochemistry. However, a scarcity of information on the polarized light-scattering properties of marine particles and the lack of a framework for separating single and multiple scattering effects on have hindered the development of polarization-based retrievals of bulk particle properties. To address these knowledge gaps, we made single scattering measurements of for several compositionally and morphologically distinct marine particle assemblages. We then used a bio-optical model to explore the influence of multiple scattering and particle characteristics on lidar measurements of made during an expedition to sample a mesoscale coccolithophore bloom. Laboratory measurements of linear depolarization revealed a complex dependency on particle shape, size, and composition that were consistent with scattering simulations for idealized nonspherical particles. Model results suggested that the variability in measured during the field expedition was driven predominantly by shifts in particle concentration rather than their bulk characteristics. However, model estimates of improved when calcite particles were represented by a distinct particle class, highlighting the influence of bulk particle properties on . To advance polarized lidar retrievals of bulk particle properties and to constrain the uncertainty in satellite lidar retrievals of particulate backscattering, these results point to the need for future efforts to characterize the variability of particulate depolarization in the ocean and to quantify the sensitivity of operational ocean lidar systems to multiple scattering.
Natural and anthropogenic controls on lake water-level decline and evaporation-to-inflow ratio in the conterminous United States
Fergus CE, Brooks JR, Kaufmann PR, Pollard AI, Mitchell R, Geldhof GJ, Hill RA, Paulsen SG, Ringold P and Weber M
Lake water levels are integral to lake function, but hydrologic changes from land and water management may alter lake fluctuations beyond natural ranges. We constructed a conceptual model of multifaceted drivers of lake water-levels and evaporation-to-inflow ratio (Evap:Inflow). Using a structural equation modeling framework, we tested our model on 1) a national subset of lakes in the conterminous United States with minimal water management to describe natural drivers of lake hydrology and 2) five ecoregional subsets of lakes to explore regional variation in water management effects. Our model fit the national and ecoregional datasets and explained up to 47% of variation in Evap:Inflow, 38% of vertical water-level decline, and 79% of horizontal water-level decline (littoral exposure). For lakes with minimal water management, Evap:Inflow was related to lake depth ( = -0.31) and surface inflow ( = -0.44); vertical decline was related to annual climate (e.g., precipitation = -0.18) and water management ( = -0.21); and horizontal decline was largely related to vertical decline ( = 0.73) and lake morphometry (e.g., depth = -0.18). Anthropogenic effects varied by ecoregion and likely reflect differences in regional water management and climate. In the West, water management indicators were related to greater vertical decline ( = 0.38), whereas in the Midwest, these indicators were related to more stable and full lake levels ( = -0.22) even during drought conditions. National analyses show how human water use interacts with regional climate resulting in contrasting impacts to lake hydrologic variation in the US.
Spatial and temporal heterogeneity of methane ebullition in lowland headwater streams and the impact on sampling design
Robison AL, Wollheim WM, Turek B, Bova C, Snay C and Varner RK
Headwater streams are known sources of methane (CH) to the atmosphere, but their contribution to global scale budgets remains poorly constrained. While efforts have been made to better understand diffusive fluxes of CH in streams, much less attention has been paid to ebullitive fluxes. We examine the temporal and spatial heterogeneity of CH ebullition from four lowland headwater streams in the temperate northeastern United States over a 2-yr period. Ebullition was observed in all monitored streams with an overall mean rate of 1.00 ± 0.23 mmol CH m d, ranging from 0.01 to 1.79 to mmol CH m d across streams. At biweekly timescales, rates of ebullition tended to increase with temperature. We observed a high degree of spatial heterogeneity in CH ebullition within and across streams. Yet, catchment land use was not a simple predictor of this heterogeneity, and instead patches scale variability weakly explained by water depth and sediment organic matter content and quality. Overall, our results support the prevalence of CH ebullition from streams and high levels of variability characteristic of this process. Our findings also highlight the need for robust temporal and spatial sampling of ebullition in lotic ecosystems to account for this high level of heterogeneity, where multiple sampling locations and times are necessary to accurately represent the mean rate of flux in a stream. The heterogeneity observed likely indicates a complex set of drivers affect CH ebullition from streams which must be considered when upscaling site measurements to larger spatial scales.
Decreasing pH impairs sexual reproduction in a Mediterranean coral transplanted at a CO vent
Marchini C, Gizzi F, Pondrelli T, Moreddu L, Marisaldi L, Montori F, Lazzari V, Airi V, Caroselli E, Prada F, Falini G, Dubinsky Z and Goffredo S
Ocean acidification, due to the increase of carbon dioxide (CO) concentration in the atmosphere and its absorption by the oceans, affects many aspects of marine calcifying organisms' biology, including reproduction. Most of the available studies on low pH effects on coral reproduction have been conducted on tropical species under controlled conditions, while little information is reported for either tropical or temperate species in the field. This study describes the influence of decreasing pH on sexual reproduction of the temperate non-zooxanthellate colonial scleractinian , transplanted in four sites along a natural pH gradient at the underwater volcanic crater of Panarea Island (Tyrrhenian Sea, Italy). The average pH values of each site (range: pH 8.07-7.40) match different scenarios of the Intergovernmental Panel on Climate Change (IPCC) for the end of the century. After 3 months under experimental conditions, the reproductive parameters of both oocytes and spermaries (abundance, gonadal index, and diameters) seem to be unaffected by low pH. However, a delay in spermary development in the pre-fertilization period and a persistence of mature oocytes in the fertilization period were observed in the most acidic site. Furthermore, no embryos were found in colonies from the two most acidic sites, suggesting a delay or an interruption of the fertilization process due to acidified conditions. These findings suggest a negative effect of low pH on sexual reproduction. However, long-term experiments, including the synergistic impact of pH and temperature, are needed to predict if this species will be able to adapt to climate change over the next century.
Coping with darkness: The adaptive response of marine picocyanobacteria to repeated light energy deprivation
Coe A, Biller SJ, Thomas E, Boulias K, Bliem C, Arellano A, Dooley K, Rasmussen AN, LeGault K, O'Keefe TJ, Stover S, Greer EL and Chisholm SW
The picocyanobacteria and are found throughout the ocean's euphotic zone, where the daily light:dark cycle drives their physiology. Periodic deep mixing events can, however, move cells below this region, depriving them of light for extended periods of time. Here, we demonstrate that members of these genera can adapt to tolerate repeated periods of light energy deprivation. Strains kept in the dark for 3 d and then returned to the light initially required 18-26 d to resume growth, but after multiple rounds of dark exposure they began to regrow after only 1-2 d. This dark-tolerant phenotype was stable and heritable; some cultures retained the trait for over 132 generations even when grown in a standard 13:11 light:dark cycle. We found no genetic differences between the dark-tolerant and parental strains of NATL2A, indicating that an epigenetic change is likely responsible for the adaptation. To begin to explore this possibility, we asked whether DNA methylation-one potential mechanism mediating epigenetic inheritance in bacteria-occurs in . LC-MS/MS analysis showed that while DNA methylations, including 6 mA and 5 mC, are found in some other strains, there were no methylations detected in either the parental or dark-tolerant NATL2A strains. These findings suggest that utilizes a yet-to-be-determined epigenetic mechanism to adapt to the stress of extended light energy deprivation, and highlights phenotypic heterogeneity as an additional dimension of diversity.
Plankton respiration in the Atacama Trench region: Implications for particulate organic carbon flux into the hadal realm
Fernández-Urruzola I, Ulloa O, Glud RN, Pinkerton MH, Schneider W, Wenzhöfer F and Escribano R
Respiration is a key process in the cycling of particulate matter and, therefore, an important control mechanism of carbon export to the ocean's interior. Most of the fixed carbon is lost in the upper ocean, and only a minor amount of organic material sustains life in the deep-sea. Conditions are particularly extreme in hadal trenches, and yet they host active biological communities. The source of organic carbon that supports them and the contribution of these communities to the ocean carbon cycle, however, remain uncertain. Here we report on size-fractionated depth profiles of plankton respiration assessed from the activity of the electron transport system in the Atacama Trench region, and provide estimates of the minimum carbon flux (C) needed to sustain the respiratory requirements from the ocean surface to hadal waters of the trench and shallower nearby sites. Plankton < 100 m contributed about 90% to total community respiration, whose magnitude was highly correlated with surface productivity. Remineralization rates were highest in the euphotic zone and declined sharply within intermediate oxygen-depleted waters, remaining fairly constant toward the bottom. Integrated respiration in ultra-deep waters (> 1000 m) was comparable to that found in upper layers, with 1.3 ± 0.4 mmol C m d being respired in the hadopelagic. The comparison between our C models and estimates of sinking particle flux revealed a carbon imbalance through the mesopelagic that was paradoxically reduced at greater depths. We argue that large fast-sinking particles originated in the overlying surface ocean may effectively sustain the respiratory carbon demands in this ultra-deep marine environment.
Carbonate fluxes by coccolithophore species between NW Africa and the Caribbean: Implications for the biological carbon pump
Guerreiro CV, Baumann KH, Brummer GA, Valente A, Fischer G, Ziveri P, Brotas V and Stuut JW
Coccolithophores are among the most important calcifying pelagic organisms. To assess how coccolithophore species with different coccolith-carbonate mass and distinct ecological resilience to ocean warming will influence the "rain ratio" and the "biological carbon pump", 1 yr of species-specific coccolith-carbonate export fluxes were quantified using sediment traps moored at four sites between NW Africa and the Caribbean (i.e., CB-20°N/21°W, at 1214 m; M1-12°N/23°W, at 1150 m; M2-14°N/37°W, at 1235 m; M4-12°N/49°W, at 1130 m). Highest coccolith-CaCO fluxes at the westernmost site M4, where the nutricline is deepest along the tropical North Atlantic, were dominated by deep-dwelling small-sized coccolith species and . Total coccolith-CaCO fluxes of 371 mg m yr at M4 were followed by 165 mg m yr at the north-easternmost CB, 130 mg m yr at M1, and 114 mg m yr at M2 in between. Coccoliths accounted for nearly half of the total carbonate flux at M4 (45%), much higher compared to 23% at M2 and 15% at M1 and CB. At site M4, highest ratios of coccolith-CaCO to particulate organic carbon fluxes and weak correlations between the carbonate of deep-dwelling species and particulate organic carbon suggest that increasing productivity in the lower photic zone in response to ocean warming might enhance the and reduce the coccolith-ballasting efficiency. The resulting weakened biological carbon pump could, however, be counterbalanced by increasing frequency of Saharan dust outbreaks across the tropical Atlantic, providing mineral ballast as well as nutrients to fuel fast-blooming and ballast-efficient coccolithophore species.
Stoichiometric imbalances complicate prediction of phytoplankton biomass in U.S. lakes: implications for nutrient criteria
Moon DL, Scott JT and Johnson TR
Using National Lakes Assessment data, we evaluated the influence of total N (TN), total P (TP), and other variables on lake chlorophyll-a concentrations. With simple linear regressions, high TN/TP samples biased predictions based on TN, and low TN/TP samples biased predictions based on TP. The bias problem was corrected, and correlation was improved, by splitting the dataset at the TN/TP ratio we estimated to be indicative of a balanced supply and developing separate regressions that predict chlorophyll-a based on TP, TN, dissolved inorganic N (DIN), dissolved organic carbon (DOC), non-algal light attenuation, depth, area, latitude, elevation, and conductivity. Both nutrients were excellent predictors, and non-algal light attenuation was the next most influential predictor. The regression analysis suggested that a potential for P only limitation (high TN/TP, 17% of samples) or N only limitation (low TN/TP, 14% of samples) can be inferred at the extremes of the TN/TP range. However, 69% of samples had an intermediate TN/TP ratio where it is difficult to infer anything about potential nutrient limitations (biomass could be N limited, P limited, N and P co-limited, or not limited by nutrients at all). Our results show that when developing phytoplankton response relationships using cross-lake datasets that span a wide range of trophic states, it is important to consider whether and how biomass is influenced by confounding factors - such as differences in the relative supply of N and P - so that biomass is not underestimated or overestimated, and nutrient criteria are not under-protective or over-protective.
Respiration by "marine snow" at high hydrostatic pressure: Insights from continuous oxygen measurements in a rotating pressure tank
Stief P, Elvert M and Glud RN
It is generally anticipated that particulate organic carbon (POC) for most part is degraded by attached microorganisms during the descent of "marine snow" aggregates toward the deep sea. There is, however, increasing evidence that fresh aggregates can reach great depth and sustain relatively high biological activity in the deep sea. Using a novel high-pressure setup, we tested the hypothesis that increasing levels of hydrostatic pressure inhibit POC degradation in aggregates rapidly sinking to the ocean interior. Respiration activity, a proxy for POC degradation, was measured directly and continuously at up to 100 MPa (corresponding to 10 km water depth) in a rotating pressure tank that keeps the aggregates in a sinking mode. Model diatom-bacteria aggregates, cultures of the aggregate-forming diatom , and seawater microbial communities devoid of diatoms showed incomplete and complete inhibition of respiration activity when exposed to pressure levels of 10-50 and 60-100 MPa, respectively. This implies reduced POC degradation and hence enhanced POC export to hadal trenches through fast-sinking, pressure-exposed aggregates. Notably, respiration measurements at ≥50 MPa revealed curved instead of linear oxygen time series whenever was present, which was not captured by respiration measurements. These curvatures correspond to alternating phases of high and low respiration activity likely connected to pressure effects on unidentified metabolic processes in .