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Microbial Iron limitation in the ocean’s twilight zone

Posted by mmaheigan

· Monday, March 31st, 2025

How deep in the ocean do microbes feel the effects of nutrient limitation? Microbial production in one third of the surface ocean is limited by the essential micronutrient iron (Fe). This limitation extends to at least the bottom of the euphotic zone, but what happens below that?

In a study that recently published in Nature we investigated the abundance and distribution of siderophores, small metabolites synthesized by bacteria to promote Fe uptake. When environmental Fe concentrations become limiting and microbes become Fe deficient, some bacteria release siderophores into the environment to bind iron and facilitate its uptake. Siderophores are therefore a window into how microbes “see” environmental Fe. We found that siderophore concentrations were high in low Fe surface waters, but surprisingly we also found siderophores to be abundant in the twilight zone (200-500 m) underlying the North and South Pacific subtropical gyres, two key ecosystems for the marine carbon cycle. In shipboard experiments with siderophores labeled with the rare ⁵⁷Fe isotope, we found rapid uptake of the label in twilight zone samples. After removing ⁵⁷Fe from the ⁵⁷Fe-siderophores complex, bacteria released the now unlabeled siderophores back into seawater to complex additional Fe (Figure. 1).

Figure 1: Iron-siderophore cycling in the twilight zone. When the seawater becomes Fe-deficient, some bacteria are able to synthesize siderophores and release them into the environment (middle left). These metabolites bind Fe (middle right) and the Fe-siderophore complex is taken up by bacteria using specialized TonB dependent transporters (TBDT; bottom right). Inside the cell, Fe is recovered from the Fe-siderophore complex (bottom left) and the siderophore excreted back into the environment to start the cycle anew.

Our results show that in large parts of the ocean microbes feel the effects of nutrient limitation deep in the water column, to at least 500 m. This greatly expands the region of the ocean where nutrients limit microbial metabolism. The effects of limitation this deep in the water column are unexplored, but twilight zone Fe deficiency could have unanticipated consequences for the efficiency of the ocean’s biological carbon pump.

Authors
Jingxuan Li, Lydia Babcock-Adams and Daniel Repeta
(all at Woods Hole Oceanographic Institution)

How do ocean microbes share the job of denitrification?

Posted by mmaheigan

· Monday, March 31st, 2025

Denitrification is a crucial multi-step process for ecosystem productivity and sustainability because some of its steps can result in the loss of the essential nutrient nitrogen or the production of greenhouse gas nitrous oxide. We do not understand why microbial functional groups conducting different steps of denitrification can coexist in the ocean and why certain groups are more abundant than others.

In a recent study published in PNAS, we uncover ecological mechanisms that govern the coexistence of these microbes. For the microbial groups utilizing different nitrogen substrates, the “stronger” groups rely on the “weaker” groups to feed them nitrogen (with respect to the organic substrates that they compete for), enabling them to coexist. For the groups competing for the same nitrogen substrates, microbes that invest more to build longer denitrification steps win the competition when nitrogen is limiting, but lose the game when nitrogen is repleted and organic carbon is limiting. The spatial and temporal variability of nutrients in the ocean allows these microbes to be observed in the same water mass.

Figure caption: Temporal and spatial heterogeneity in nutrients promotes the coexistence of functionally diverse denitrifiers in the ocean.

These hypothesized coexistence patterns help us predict where and when nitrogen loss and nitrous oxide production may occur. As human activities continue to alter marine nutrient balances, these predictions help us better anticipate ocean responses and design better strategies for mitigating negative anthropogenic impacts on the ocean.

Authors
Xin Sun (Carnegie Institution for Science) @xinsun-putiger.bsky.social
Emily Zakem (Carnegie Institution for Science) @carnegiescience.bsky.social

Tiny parasites, big impact: Species networks and carbon recycling in an oligotrophic ocean

Posted by mmaheigan

· Tuesday, March 12th, 2024

Parasites are everywhere in the ocean. Including the microbial realm where a diverse, widespread group of protist parasites (Syndiniales) infect and kill a range of hosts, such as dinoflagellates, radiolarians, and even larger zooplankton. A complete Syndiniales infection cycle is only 2-3 days. First, the parasite is a free-living spore. Once inside a host, the parasite consumes the host’s carbon and becomes a larger multicellular organism (a trophont) eventually causing the host to burst open and release hundreds of new spores.

Like viruses, parasite lysis is expected to reroute organic carbon to the microbial loop, potentially decreasing the amount of carbon available for export to the deep sea. Yet, the role of Syndiniales in carbon cycling has been hard to define, as depth-specific infection dynamics and links to carbon export remain poorly understood.

Parasites are everywhere in the ocean. Including the microbial realm where a diverse, widespread group of protist parasites (Syndiniales) infect and kill a range of hosts, such as dinoflagellates, radiolarians, and even larger zooplankton. A complete Syndiniales infection cycle is only 2-3 days. First, the parasite is a free-living spore. Once inside a host, the parasite consumes the host’s carbon and becomes a larger multicellular organism (a trophont) eventually causing the host to burst open and release hundreds of new spores.

Figure 1. The mean relative abundance of Syndiniales (purple) in the photic zone (<140 m) is negatively correlated with particulate organic carbon (POC) flux at 150 m (p-value < 0.001). Similar correlations are not significant (p-values > 0.05) for other major 18S taxonomic groups, like Dinophyceae (red) and Arthropoda (green).

In a recent study published in ISME Communications, authors analyzed an 18S rRNA gene metabarcoding dataset from the Bermuda Atlantic Time-series Study (BATS) site that included 4 years (2016-2019) and twelve depths (1-1000 m). Syndiniales were the most dominant 18S group at BATS, present throughout the photic and aphotic zones. These parasites were prominent in species networks constructed with 18S sequence data, with significant associations with dinoflagellates and copepods in the surface, and with radiolarians in the aphotic zone. In addition, Syndiniales were the only major 18S group to be significantly (and negatively) correlated to particulate carbon flux (at 150 m), which was estimated from sediment trap data collected concurrently at BATS (Figure 1). This is in situ evidence of flux attenuation among Syndiniales, as they recycle host carbon that would otherwise transfer up to larger organisms (e.g., via grazing). Lastly, authors found 19% of the Syndiniales community is linked between photic and aphotic zones, indicating that parasites are sinking on particles and/or are recirculated via diel vertical migration. Overall, these findings elevate the role of Syndiniales in microbial food webs and further emphasize the importance in quantifying parasite-host dynamics to inform ocean carbon models.

Authors
Sean Anderson (University of New Hampshire / Woods Hole Oceanographic Institution)
Leocadio Blanco-Bercial (Bermuda Institute of Ocean Sciences / Arizona State University)
Craig Carlson (University of California, Santa Barbara)
Elizabeth Harvey (University of New Hampshire)

Identifying the water mass composition of a sample has never been so easy!

Posted by mmaheigan

· Thursday, August 31st, 2023

When we collect seawater in any point of the ocean, we are collecting a mix of water masses from different origin that traveled until there keeping their salinity and temperature properties. The Atlantic Ocean is likely the most complex basin in term of water masses containing more than 15 in its depths. Some of them were “born” in the North Atlantic Ocean, others in the Southern Ocean, even in the Mediterranean Sea! And when we collect a seawater sample we can know which water masses are there, where they come from, what happened to each of them during their journey to us, what story can they tell us.

The variation of any non-conservative property (such as dissolved organic carbon or nutrients) in the deep open ocean depends on the mixing of those water masses and on the biogeochemical processes affecting it (such as heterotrophic respiration). But the effect of the water mass mixing is usually very high, so in order to study the biogeochemical processes, it is necessary to remove that effect.

On the other hand, estimating the contribution of the water masses composing a sample is useful to trace the distribution of each water mass identifying the depth of maximum water mass contribution or the depth-range where the water mass is dominant contributing > 50%. Ocean biogeochemists and microbiologists can get more out of their data estimating the impact of water mass mixing on the variability of any chemical (e.g. inorganic nutrients and dissolved organic carbon) or biological (e.g. prokaryotic heterotrophic abundance and production) property.

Knowing the contribution of each water mass to each sample was not an easy task and required expertise on the origin, circulation and mixing patterns of the water masses present in the study area. This could be even harder in very complex oceanic basin such as the deep Atlantic Ocean. The most commonly used methodology is the Optimum Multi-Parameter (OMP) analysis that was first applied by Tomczak in 1981. However, this methodology is time consuming and requires availability of a large set of quality-controlled chemical variables (e.g. nutrients, oxygen,..) together with a deep knowledge of the oceanography of the studied area. Those chemical variables are not always available or do not have the required quality by contrast to potential temperature and salinity that are high standard core variables in any cruise or database. In a recent research article, we applied multi-regression machine learning models to solve ocean water mass mixing. The models tested were trained using the solutions from OMP analyses previously applied to samples from cruises in the Atlantic Ocean. Extremely Randomized Trees algorithm yielded the highest score (R² = 0.9931; mse = 0.000227). The model allows solving the mixing of water masses in the Atlantic Ocean using potential temperature, salinity, latitude, longitude and depth. Potential temperature and salinity are the most commonly collected and curated variables in oceanography both from oceanographic cruises and autonomous vehicles (e.g. ARGO) avoiding the use of less commonly measured chemical variables which also require longer and time-consuming analyses of both the water samples and the data.

Figure 1. A16 section for the contribution of the water masses (A) AAIW5, (B) ENACW12, (C) AAIW3, (D) MW, (E) LSW, (F) ISOW, (G) CDW and (H) WSDW obtained with the Extremely Randomized Trees algorithm. Ocean Data View software (Schlitzer, 2015).

We also provide the code with instructions where any user can easily introduce the required variables (latitude, longitude, depth, temperature and salinity) of the chosen Atlantic samples and obtain the water mass proportion of each one in a fast and easy way. Actually, it would allow the user to obtain this information in real time during a cruise.

New research using other methods like OMP and its variants can be incorporated to the existing model increasing its accuracy and prediction capacity. Help us to improve the model and increase its spatial resolution!

Ocean biogeochemists and microbiologists can benefit from this tool even if they do not have a deep knowledge of the oceanography of the studied area. Identifying the water masses composition of a sample has never been so easy!

Author
Cristina Romera-Castillo (Instituto de Ciencias del Mar-CSIC, Barcelona, Spain)

Twitter: @crisrcas

Hydrostatic pressure substantially reduces deep-sea microbial activity

Posted by mmaheigan

· Thursday, May 11th, 2023

Deep sea microbial communities are experiencing increasing hydrostatic pressure with depth. It is known that some deep sea microbes require high hydrostatic pressure for growth, but most measurements of deep-sea microbial activity have been performed under atmospheric pressure conditions.

In a recent paper published in Nature Geoscience, the authors used a new device coined ‘In Situ Microbial Incubator’ (ISMI) to determine prokaryotic heterotrophic activity under in situ conditions. They compared microbial activity in situ with activity under atmospheric pressure at 27 stations from 175 to 4000 m depths in the Atlantic, Pacific, and the Southern Ocean. The bulk of heterotrophic activity under in situ pressure is always lower than under atmospheric pressure conditions and is increasingly inhibited with increasing hydrostatic pressure. Single-cell analysis revealed that deep sea prokaryotic communities consist of a small fraction of pressure-loving (piezophilic) microbes while the vast majority is pressure-insensitive (piezotolerant). Surprisingly, the piezosensitive fraction (~10% of the total community) responds with a more than 100-fold increase of activity upon depressurization. In the microbe proteomes, the authors uncovered taxonomically characteristic survival strategies in meso- and bathypelagic waters. These findings indicate that the overall heterotrophic microbial activity in the deep sea is substantially lower than previously assumed, which implies major impacts on the carbon budget of the ocean’s interior.

Figure caption: Deep sea microbial activity under varying pressure. (a) In situ bulk leucine incorporation rates normalized to rates obtained at atmospheric pressure conditions. (b) A microscopic view of a 2000 m sample collected in the Atlantic and incubated under atmospheric pressure conditions. The black halos around the cells are silver grains corresponding to their activities. The highly active cells (indicated by arrows) were rarely found in in situ pressure incubations. (c) Depth-related changes in the metaproteome of three abundant deep sea bacterial taxa (Alteromonas, Bacteroidetes, and SAR202). The number indicates shared and unique up- and down-regulated proteins in different depth zones.

Authors
Chie Amano (University of Vienna, Austria)
Zihao Zhao (University of Vienna, Austria)
Eva Sintes (University of Vienna, IEO-CSIC, Spain)
Thomas Reinthaler (University of Vienna, Austria)
Julia Stefanschitz (University of Vienna, Austria)
Murat Kisadur (University of Vienna, Austria)
Motoo Utsumi (University of Tsukuba, Japan)
Gerhard J. Herndl (University of Vienna, Netherlands Institute for Sea Research)

Twitter @microbialoceanW

Does dark carbon fixation supply labile DOC to the deep ocean?

Posted by mmaheigan

· Thursday, March 30th, 2023

Nitrifying microbes are the most abundant chemoautotrophs in the dark ocean. Though better known for their role in the nitrogen cycle, they also fix dissolved inorganic carbon (DIC) into biomass and thus play an important role in the global carbon cycle. The release of organic compounds by these microbes may represent an as-yet unaccounted for source of dissolved organic carbon (DOC) available to heterotrophic marine food webs. Quantifying how much DIC these microbes fix and release again into the ambient seawater is critical to a complete understanding of the carbon cycle in the deep ocean.

To address this knowledge gap, a recent study grew ten diverse nitrifier cultures and measured their cellular carbon (C) content, DIC fixation yields and DOC release rates. The results indicate that nitrifiers release between 5 and 15% of their recently fixed DIC as DOC (Figure 1). This would equate to global ocean fluxes of 0.006–0.02 Pg C yr⁻.

Figure 1. DOC release by ten different chemoautotrophic nitrifying (ammonia- and nitrite-oxidizing) microbes. The diversity of marine nitrifiers used in this study comprises all genera currently available as axenic cultures. Species and strain names are given for completeness.

Our results provide values for biogeochemical models of the global carbon cycle, and help to further constrain the relationship between C and N fluxes in the nitrification process. Elucidating the lability and fate of carbon released by nitrifiers will be the crucial next step to understand its implications for marine food-web functioning and the biological sequestration of carbon in the ocean.

Authors:
Barbara Bayer (University of California, Santa Barbara and University of Vienna)
Kelsey McBeain (University of California, Santa Barbara)
Craig A. Carlson (University of California, Santa Barbara)
Alyson E. Santoro (University of California, Santa Barbara)

Linking the calcium carbonate and alkalinity cycles in the North Pacific ocean

Posted by mmaheigan

· Tuesday, December 13th, 2022

The marine carbon and alkalinity cycles are tightly coupled. Seawater stores so much carbon because of its high alkalinity, or buffering capacity, and the main driver of alkalinity cycling is the formation and dissolution of biologically produced calcium carbonate (CaCO3). In a recent publication in GBC, the authors conducted novel carbon-13 tracer experiments to measure the dissolution rates of biologically produced CaCO3 along a transect in the North Pacific Ocean. They combined these experiment data with shipboard analyses of the dissolved carbonate system, the 13C-content of dissolved inorganic carbon, and CaCO3 fluxes, to constrain the alkalinity cycle in the upper 1000 meters of the water column. Dissolution rates were too slow to explain alkalinity production or CaCO3 loss from the particulate phase. However, driving dissolution with the metabolic consumption of oxygen brings alkalinity production and CaCO3 loss estimates into quantitative agreement (Figure). The authors argue that a majority of CaCO3 production is likely dissolved through metabolic processes in the upper ocean, including zooplankton grazing, digestion, and egestion, and microbial degradation of marine particle aggregates that contain both organic carbon and CaCO3. This hypothesis stems from the basic fact that almost all marine CaCO3 is biologically produced, placing CaCO3 at the source of the acidifying process (metabolic consumption of organic matter). This process is important because it puts an emphasis on biological processing for the cycling of not only carbon, but also alkalinity, the main buffering component in seawater. These results should help both scientists and stakeholders to understand the fundamental controls on calcium carbonate cycling in the ocean, and therefore the processes that distribute alkalinity throughout the world’s oceans.

Figure Caption: Sinking-dissolution model results compared with tracer-based alkalinity regeneration rates (TA*-CFC, Feely et al., 2002). We also plot alkalinity regeneration rates using updated time transit distribution ages (TA*- and Alk*-TTD). The modeled alkalinity regeneration rate uses our measured dissolution rates for biologically produced calcite and aragonite, and is driven by a combination of background saturation state and metabolic oxygen consumption. The dissolution rate is split up into a calcite component (produced mainly by coccolithophores) and an aragonite component (produced mainly by pteropods). Aragonite does not contribute significantly to the overall dissolution rate. Driving dissolution by metabolic oxygen consumption produces alkalinity regeneration rates that are in quantitative agreement with tracer-based estimates.

Authors:
Adam Subhas (Woods Hole Oceanographic Institution) et al.

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Marine heatwave implications for future phytoplankton blooms

Posted by mmaheigan

· Thursday, October 15th, 2020

Ocean temperature extreme events such as marine heatwaves are expected to intensify in coming decades due to anthropogenic warming. Although the effects of marine heatwaves on large plants and animals are becoming well documented, little is known about how these warming events will impact microbes that regulate key biogeochemical processes such as ocean carbon uptake and export, which represent important feedbacks on the global carbon cycle and climate.

Figure caption: Relationship between phytoplankton bloom response to marine heatwaves and background nitrate concentration in the 23 study regions. X-axis denotes the annual-mean sea-surface nitrate concentration based on the model simulation (1992-2014; OFAM3, blue) and the in situ climatology (WOA13, orange). Y-axis denotes the mean standardised anomalies (see Equation 1 of the paper) of simulated sea-surface phytoplankton nitrogen biomass (1992-2014; OFAM3, blue) and observed sea-surface chlorophyll a concentration (2002-2018; MODIS, orange) during the co-occurrence of phytoplankton blooms and marine heatwaves.

In a recent study published in Global Change Biology, authors combined model simulations and satellite observations in tropical and temperate oceanographic regions over recent decades to characterize marine heatwave impacts on phytoplankton blooms. The results reveal regionally‐coherent anomalies depicted by shallower surface mixed layers and lower surface nitrate concentrations during marine heatwaves, which counteract known light and nutrient limitation effects on phytoplankton growth, respectively (Figure 1). Consequently, phytoplankton bloom responses are mixed, but derive from the background nutrient conditions of a study region such that blooms are weaker (stronger) during marine heatwaves in nutrient-poor (nutrient-rich) waters.

Given the projected expansion of nutrient-poor waters in the 21st century ocean, the coming decades are likely to see an increased occurrence of weaker blooms during marine heatwaves, with implications for higher trophic levels and biogeochemical cycling of key elements.

Authors:
Hakase Hayashida (University of Tasmania)
Richard Matear (CSIRO)
Pete Strutton (University of Tasmania)

Will global change “stress out” ocean DOC cycling?

Posted by mmaheigan

· Tuesday, September 29th, 2020

The dissolved organic carbon (DOC) pool is vital for the functioning of marine ecosystems. DOC fuels marine food webs and is a cornerstone of the earth’s carbon cycle. As one of the largest pools of organic matter on the planet, disruptions to marine DOC cycling driven by climate and environmental global changes can impact air-sea CO2 exchange, with the added potential for feedbacks on Earth’s climate system.

Figure 1. Simplified view of major dissolved organic carbon (DOC) sources (black text) and sinks (yellow text) in the ocean.

Since DOC cycling involves multiple processes acting concurrently over a range of time and space scales, it is especially challenging to characterize and quantify the influence of global change. In a recent review paper published in Frontiers in Marine Science, the authors synthesize impacts of global change-related stressors on DOC cycling such as ocean warming, stratification, acidification, deoxygenation, glacial and sea ice melting, inflow from rivers, ocean circulation and upwelling, and atmospheric deposition. While ocean warming and acidification are projected to stimulate DOC production and degradation, in most regions, the outcomes for other key climate stressors are less clear, with much more regional variation. This synthesis helps advance our understanding of how global change will affect the DOC pool in the future ocean, but also highlights important research gaps that need to be explored. These gaps include for example a need for studies that allow to understand the adaptation of degradation/production pathways to global change stressors, and their cumulative impacts (e.g. temperature with acidification).

Authors:
C. Lønborg (Aarhus University)
C. Carreira (CESAM, Universidade de Aveiro)
Tim Jickells (University of East Anglia)
X.A. Álvarez-Salgado (CSIC, Instituto de Investigacións Mariñas)

Blue hole in the South China Sea reveals ancient carbon

Posted by mmaheigan

· Wednesday, July 8th, 2020

Blue holes are unique depositional environments that are formed within carbonate platforms. Due to an enclosed geomorphology that restricts water exchange, blue hole ecosystems are typically characterized by steep biogeochemical gradients and distinctive microbial communities. For the past three decades, studies have described vertical gradients in physical, chemical, and biological parameters that typify blue hole water columns, but their elemental cycles, particularly carbon, remain poorly understood.

Figure 1. Aerial photo of the Yongle Blue Hole in the South China Sea (Credit: P. Yao et al./JGR Biogeosciences)

In July 2016, the Yongle Blue Hole (YBH) was discovered to be the deepest known blue hole on Earth (~300 m). YBH is located in the Xisha Islands of the South China Sea. The unique features and ease of accessibility make YBH an ideal natural laboratory for studying carbon cycling in marine anoxic systems. In a recent study published in JGR Biogeosciences, the authors reported extremely low concentrations of dissolved organic carbon (DOC) (e.g., 22 µM) and very high concentrations of dissolved inorganic carbon (DIC) (e.g., 3,090 µM) in YBH deep waters. Radiocarbon dating revealed that the YBH DOC and DIC were unusually old, yielding ages (6,810 and 8270 years BP, respectively) that are much more typical of open ocean deep water. Based on H₂S and microbial community composition profiles, the authors concluded that sharp redox gradients and a high abundance of sulfur cycling bacteria were likely responsible for much of the DOC consumption in YBH. The unusually low concentrations and old DOC ages in the relatively shallow YBH suggest short-term cycling of recalcitrant DOC in oceanic waters, which has been recognized as a long-term microbial carbon sink in the global ocean. The stoichiometry of DIC and total alkalinity changes suggested that the accumulation of DIC in the deep layer of the YBH was largely derived from both the dissolution of carbonate and OC decomposition through sulfate reduction. However, the role of carbonate dissolution from the walls of the blue hole in affecting the old ages of carbon in this system remain uncertain, yet there appears to no evidence of subterranean freshwater into the bottom waters of the blue hole. In the face of expanding oxygen minimum zones and anthropogenically-induced coastal hypoxia, blue holes such as YBH can provide an accessible natural laboratory in which to study the microbial and biogeochemical features that typify these low-oxygen systems.

Authors:
Peng Yao (Ocean University of China)
Thomas S. Bianchi (University of Florida)
Xuchen Wang (Ocean University of China)
Zuosheng Yang (Ocean University of China)
Zhigang Yu (Ocean University of China)

Funding for the Ocean Carbon & Biogeochemistry Project Office is provided by the National Science Foundation (NSF) and the National Aeronautics and Space Administration (NASA). The OCB Project Office is housed at the Woods Hole Oceanographic Institution.

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