Biology subjects

Carlson, C. A.

Publications and source records attributed to Carlson, C. A..

14 recordsLinked to original sources

bioRxiv2026

Particle lability drives degradation dynamics and bacterial community assembly during a Phaeocystis bloom decline

Microbial degradation of suspended and sinking organic carbon regulates long-term oceanic carbon storage by controlling the efficiency of the biological pump. Yet microbial controls on carbon export and remineralization remain poorly constrained, limiting predictions of how ocean carbon cycling will respond to climate change. Here, we combined in situ sampling with ship-based incubations to quantify prokaryote-driven removal rates of suspended and sinking total organic carbon (TOC). Samples were collected below the mixed layer during three stages of a spring Phaeocystis pouchetii bloom in the Labrador Sea. Phaeocystis blooms can dominate regional phytoplankton biomass and are expected to increase under future climate. Removal rates were used as a proxy for carbon lability and combined with 16S rRNA metabarcoding and carbon composition analyses to link microbial community structure with substrate characteristics. Removal rates of sinking particles (0.02-0.06 d-1) were an order of magnitude higher than those of suspended TOC (0.002 d-1) during bloom-decline and non-bloom. In contrast, during late-bloom, suspended carbon exhibited rates of 0.01 d-1, comparable to sinking particles, and was enriched in exopolymer-rich colonies. Prokaryotic community composition varied primarily among bloom stages rather than carbon fractions, indicating that bloom stage-- and thus particle origin and composition--was the dominant control on bacterial degradation and assembly. Bacterial diversity peaked where carbon was refractory and originated from mixed phytoplankton. Together, these results demonstrate that suspended Phaeocystis-derived carbon can be rapidly remineralized when blooms produce exopolymer-rich colonies and highlight bloom stage as key regulator of microbial carbon processing and biological pump efficiency.

microbiology↗

bioRxiv2026

Iron limitation alters diatom carbon flow through shifts in microbiome exometabolite consumption

Iron is required for photosynthesis, and thus affects biogeochemical cycling in widespread regions where its availability is limited. Turnover of aquatic photosynthetically-derived carbon is largely constrained by bacterial activity, but we lack a mechanistic understanding of how iron limitation influences this activity. We examined a bacterial enrichment community dependent on carbon from the diatom Phaeodactylum tricornutum to investigate how iron limitation alters the flow of carbon to bacteria, and exometabolite and community composition. Using stable isotope tracing, we quantified diatom exudate incorporation with single-cell-resolution. We identified a population of bacteria under iron limitation with high metabolic activity yet low incorporation of newly-fixed diatom carbon, indicating a shift in metabolism relative to the iron-replete control. Ultra-high-resolution exometabolomics revealed bacterial consumption of aromatics, lipid-like compounds, and purines and pyrimidines occurred under iron-limitation, when these compounds also exhibited increased exudation. We identified gene pathways for utilization of these compounds in taxa with increased abundance under iron limitation which may be responsible for carbon flow shifts. These results provide a mechanistic link between iron-limitation driven shifts in exudate composition and flow of carbon to the microbiome. This has important implications for predicting carbon flow in surface oceans and manipulating algal-bacterial interactions in engineered systems.

microbiology↗

bioRxiv2025

Seasonal and developmental stage changes in exudate carbohydrate content shape the kelp microbiome

Photoautotrophs release a large amount of their fixed carbon as exudates, which shapes their immediate environment, including the composition of their microbiome. Here we evaluated the microbiome and exudate composition of Macrocystis pyrifera (giant kelp), a globally distributed foundation species, in response to seasonal nutrient availability and developmental stage. We combine 16S rRNA amplicon analysis of the giant kelp microbiome with carbohydrate monomer analysis of kelp exudates to examine microbe-exudate relationships. We found significant differences in the microbiome and exudate composition between seasons and developmental stages of giant kelp. Higher tissue-nitrogen content in the spring coincided with elevated amounts of glucosamine, a nitrogen-containing sugar, in giant kelp exudates, while senescence led to the shedding of mannuronic acid, an alginate indicator. The release of glucosamine and fucose-rich exudates was correlated with an increase in the relative abundance of bacteria within the Planctomycetota phylum, whereas mannuronic acid-rich exudates coincided with an increase in the relative abundance of members of the Flavobacteria and Gammaproteobacteria lineages. We investigated putative carbohydrate-microbe relationships by isolating a member of the Planctomycetota phylum from the surface of giant kelp. We demonstrate that the growth of this isolate on fucoidan and N-acetyl glucosamine, but not alginate reflects the observed relative abundance of this clade in the kelp microbiome in response to variable carbohydrate exudation. This suggests a key role of kelp mucilage carbohydrate composition in structuring its microbiome as has been observed for other organisms such as corals and within the human gut.

ecology↗

bioRxiv2025

Characterizing organisms from three domains of life with universal primers from throughout the global ocean

We introduce the Global rRNA Universal Metabarcoding Plankton database (GRUMP), which consists of 1194 samples that were collected from 2003-2020 and cover extensive latitudinal and longitudinal transects, as well as depth profiles in all major ocean basins. DNA from unfractionated (> 0.2{micro}m) seawater samples was amplified using the 515Y/926R universal three- domain rRNA gene primers, simultaneously quantifying the relative abundance of amplicon sequencing variants (ASVs) from bacteria, archaea, eukaryotic nuclear 18S, and eukaryotic plastid 16S. Thus, the ratio between taxa in one sample is directly comparable to the ratio in any other GRUMP sample, regardless of gene copy number differences. This obviates a problem in prior global studies that used size-fractionation and different rRNA gene primers for bacteria, archaea, and eukaryotes, precluding comparisons across size fractions or domains. On average, bacteria contributed 71%, eukaryotes 19%, and archaea 8% to rRNA gene abundance, though eukaryotes contributed 32% at latitudes > 40{degrees}. GRUMP is publicly available on the Simons Collaborative Marine Atlas Project (CMAP), promoting the global comparison of marine microbial dynamics.

ecology↗

bioRxiv2024

Contribution of ammonia oxidizers to inorganic carbon fixation in the dark ocean

Ammonia-oxidizing archaea are the most abundant chemolithoautotrophs in the ocean, comprising up to 40% of microbial cells in deep waters, and are assumed to dominate dissolved inorganic carbon (DIC) fixation below the sunlit surface layer. Yet, the supply of reduced nitrogen from particulate organic matter flux from the surface is insufficient to support the amount of nitrification required to sustain measured DIC fixation rates in the dark ocean. The aim of this study was to quantify the contribution of ammonia oxidizers to DIC fixation in the dark ocean. We used phenylacetylene - a specific inhibitor of the ammonia monooxygenase enzyme - to selectively inhibit ammonia oxidizers during two oceanographic expeditions in the eastern tropical and subtropical Pacific Ocean spanning 35{o} N to 10{o} S. We show that ammonia oxidizers contribute only a small fraction to dark DIC fixation, accounting for 2 to 22% of the depth-integrated rates in the eastern tropical Pacific. The highest contributions were observed at the depth of the nitrification maximum, where ammonia oxidation could account for up to 50% of dark DIC fixation. Our results help to reconcile the observed discrepancies between nitrogen supply and DIC fixation at depth, and provide a new perspective on global ocean chemolithoautotrophy, revealing that the majority of DIC fixation within the lower euphotic zone and below 200 m depth is not fueled by ammonia oxidation. SignificanceMicrobes in the ocean play important roles in the global carbon cycle and the oceans capacity to sequester carbon. Despite this importance, deciphering the contributions of different microbial metabolic processes to the oceanic carbon budget remains challenging. Particularly in the dark ocean, large discrepancies between organic matter fluxes and measured microbial metabolic rates are observed. We show that abundant chemoautotrophs - ammonia-oxidizing microbes - contribute only a small fraction to dark carbon fixation in the Pacific Ocean, challenging the current view that carbon fixation in the dark ocean is primarily sustained by nitrification. This work advances our understanding of microbial carbon processing, and offers new insights into the long-standing question of the main energy sources fueling carbon fixation in the dark ocean.

microbiology↗

bioRxiv2024

Seasonal exometabolites are regulated by essential microbial metabolisms in the oligotrophic ocean

Predictions of how the biogeochemical reservoir of marine dissolved organic matter (DOM) will respond to future ocean changes require an improved understanding of the thousands of individual microbe-molecule interactions which regulate the transformation and fate of DOM. Bulk characterizations of organic matter can mask this complex network of interactions comprised of rich chemical and taxonomic diversity. Here, we present a three-year, depth-resolved time-series of the seasonal dynamics of the exometabolome and the bacterioplankton community at the Bermuda Atlantic Time-series Study (BATS) site. We find both time-series to be highly structured and compositionally distinct across sampling depths. Putative exometabolite identifications (gonyol, glucose 6-sulfate, succinate, and trehalose) indicate that at least a portion of the exometabolome contains rapidly remineralized, labile molecules. We hypothesize that apparent seasonal accumulation of these labile molecules could result from environmental conditions that alter community composition on a seasonal timescale and thus shift the relative proportions of microbial functions that produce and consume the substrates. Critically, we found the composition of seasonal DOM features was more stable interannually than the microbial community structure. By estimating redundancy of metabolic functions responsible for cycling these molecules in BATS metagenomes, we propose a paradigm whereby core microbial metabolisms, either those utilized by all or by a subset of marine microbes, are better predictors of DOM composition than microbial taxonomies. The molecular-level characterization of DOM achieved herein highlights the metabolic imprint of microbial activity in DOM composition and greatly enhances our understanding of the dynamics regulating Earths largest reservoir of organic carbon. Significance statementMarine dissolved organic matter (DOM) is a major carbon reservoir that acts as a critical control on Earths climate. DOM dynamics are largely regulated by a complex web of microbial interactions, but the mechanisms underpinning these processes are not well understood. In a three-year time-series, we found thousands of DOM molecules and microbial taxa exhibited seasonal patterns. Critically, the identity of the microbes was more variable between years than the composition of the DOM molecules. We suggest that shared metabolisms encoded by genes that conduct core microbial functions are responsible for the more stable composition of DOM. This work links DOM molecules with microbial biodiversity, and presents testable predictors of DOM composition in our changing oceans.

microbiology↗

bioRxiv2024

Predictable functional biogeography of marine microbial heterotrophs

Heterotrophic bacteria and archaea ( heteroprokaryotes) drive global carbon cycling, but how to quantitatively organize their functional complexity remains unclear. We generated a global-scale understanding of marine heteroprokaryotic functional biogeography by synthesizing genetic sequencing data with a mechanistic marine ecosystem model. We incorporated heteroprokaryotic diversity into the trait-based model along two axes: substrate lability and growth strategy. Using genetic sequences along three ocean transects, we compiled 21 heteroprokaryotic guilds and estimated their degree of optimization for rapid growth (copiotrophy). Data and model consistency indicated that gradients in grazing and substrate lability predominantly set biogeographical patterns, and identified deep-ocean slow copiotrophs whose ecological interactions control the surface accumulation of dissolved organic carbon.

ecology↗

bioRxiv2023

Microbial Respiration in Contrasting Ocean Provinces via High-Frequency Optical Assays

Microbial respiration plays a pivotal role in the marine carbon cycle, influencing the fraction of fixed carbon that undergoes remineralization versus export to depth. Despite its importance, methodological constraints have led to an inadequate understanding of this process, especially in low-activity oligotrophic and mesopelagic regions. Here, we quantify respiration rates as low as 0.2 {micro}mol O2 L-1 d-1 in contrasting ocean productivity provinces using optical oxygen sensors to identify size-fractionated respiration trends. At the low productivity North Pacific Ocean Station Papa, surface microbial respiration was relatively stable at 1.2 {micro}mol O2 L-1 d-1. Below the surface there was a decoupling of respiration and bacterial production potentially driven by phytodetritus remineralization. Size-fractionated rates showed cells <5 {micro}m contributed the most to Pacific respiration. At the North Atlantic Porcupine Abyssal Plain, the optode measurement frequency was drastically increased. Surface microbial respiration was higher (1.7 {micro}mol O2 L-1 d-1) and decreased by 3-fold below the euphotic zone. The Atlantic filtered fraction contributions to total respiration shifted with the phytoplankton bloom evolution. The high resolution optode method used in the Atlantic is consistent with coinciding in vivo 2-para-(iodophenyl)-3(nitrophenyl)-5(phenyl) tetrazolium chloride respiratory stain measurements and historical site estimates. We estimate that 58% of gross primary production was respired at the Pacific site and 34% at the Atlantic site, demonstrating that the Atlantic had a higher carbon export potential. This study highlights the dynamic nature of respiration across vertical, temporal, and size-fractionated factors, emphasizing the need for sensitive, high-throughput techniques to better understand ocean ecosystem metabolism.

microbiology↗

bioRxiv2023

Syndiniales parasites drive species networks and are a biomarker for carbon export in the oligotrophic ocean

Microbial associations that result in phytoplankton mortality are important for carbon transport in the ocean. This includes parasitism, which in microbial food webs, is dominated by the marine alveolate group, Syndiniales. Parasites are expected to contribute to carbon recycling via host lysis; however, knowledge on host dynamics and correlation to carbon export remain unclear and limit the inclusion of parasitism in biogeochemical models. We analyzed a 4-year 18S rRNA metabarcoding dataset (2016-2019), performing network analysis for twelve discrete depths (1- 1000 m) to determine Syndiniales-host associations in the seasonally oligotrophic Sargasso Sea. Analogous water column and sediment trap data were included to define environmental drivers of Syndiniales and their correlation with particulate carbon flux (150 m). Syndiniales accounted for 48-74% of network edges, most often associated with Dinophyceae and Arthropoda (mainly copepods) at the surface and Rhizaria (Polycystinea, Acantharea, and RAD-B) in the aphotic zone. Unlike other major groups, Syndiniales were significantly (and negatively) correlated with particulate carbon flux, suggesting parasites may drive flux attenuation through remineralization. Examination of Syndiniales amplicons revealed a range of depth patterns, including specific ecological niches and vertical connection among a subset (19%) of the community, the latter implying sinking of parasites (infected hosts or spores) on particles. Our findings point to the use of Syndiniales as biomarkers of carbon export, highlighting their importance for marine food webs and biogeochemistry. Significance StatementSyndiniales parasites are widespread in the ocean and represent a potentially important, albeit poorly resolved, source of carbon recycling. Here, we assess Syndiniales population dynamics, trophic relationships, and links to carbon export in the Sargasso Sea. Species networks at all depths were driven by Syndiniales, with parasite-host relationships varying with depth based on shifts in host composition. Syndiniales were the only eukaryote group to be significantly (and negatively) correlated with particulate carbon flux, indicating their contribution to flux attenuation via remineralization. Yet, a subset of parasites was vertically connected between photic and aphotic zones, suggesting continued export. Our findings elevate the critical role of Syndiniales in marine microbial systems and reveal their potential use as biomarkers for carbon export.

microbiology↗

bioRxiv2023

Quantitative analysis of food web dynamics in a low export ecosystem

Food webs trace the flow of organic matter and energy among producers and consumers; for pelagic marine food webs, network complexity directly influences the amount and form of carbon exported to the deep ocean via the biological pump. Here we present a synoptic view of mixed layer food web dynamics observed during the late summer 2018 EXport Processes in the Ocean from Remote Sensing (EXPORTS) field campaign in the subarctic Northeast Pacific at the long-running time-series site, Ocean Station Papa. Carbon biomass reservoirs of phytoplankton, microzooplankton, and bacterioplankton, were approximately equal while mesozooplankton biomass was 70% lower. Live organisms composed [~]40% of the total particulate organic carbon within the mixed layer: the remainder was attributed to detritus. Rates of carbon transfer among reservoirs indicated production and assimilation rates were well balanced by losses, leaving little organic carbon available for export. The slight positive net community production rate generated organic carbon that was exported from the system in the form of food web byproducts, such as large fecal pellets generated by mesozooplankton. This characteristically regenerative food web had relatively slow turnover times with small-magnitude transfers of carbon relative to standing stocks that occurred amidst a high background concentration of detrital particles and dissolved organic matter. The concurrent estimation of food web components and rates revealed that separated processes dominated the transfer of carbon within the food web compared to those that contributed to export. Plain Language SummaryThe biological carbon pump drives a downward flux of organic matter from the sunlit surface ocean to the vast ocean interior. Ecological interactions in the surface ocean directly affect the amount and type of carbon that is exported to the deep ocean. In this study, we present a synthesis of the late summer mixed layer food web in the Northeast Pacific that was extensively characterized during the 2018 EXport Processes in the Ocean from Remote Sensing (EXPORTS) field campaign. We found the majority of carbon was recycled within the mixed layer by microbes through multiple transfers between producers and consumers. Larger organisms, mesozooplankton and salps, only consumed a small amount of carbon but through the formation of sinking fecal pellets were the main mechanism of transporting carbon out of the system. The study highlights the need to concurrently study microbial and large organism dynamics to develop a predictive understanding of the fate of organic carbon in the oceans. Key PointsO_LIThe microbial loop dominated carbon flow in the late summer mixed layer food web of the North Pacific, most net production was respired leaving little carbon available for export. C_LIO_LIActive production and consumption of organic carbon occurred amid a high background of detrital particulate organic carbon (58% of total) with slow turnover time, 66 d. C_LIO_LIMesozooplankton which had relatively minor carbon consumption rates created the majority of export production due to efficient repackaging of consumed material. C_LI

ecology↗

bioRxiv2022

Seasonal and daily patterns in known dissolved metabolites in the northwestern Sargasso Sea

Organic carbon in seawater plays a significant role in the global carbon cycle. The concentration and composition of dissolved organic carbon, operationally defined in this project as organic carbon that passes through a 0.2 {micro}m filter, reflect the activity of the biological community and chemical reactions that occur in seawater. From 2016 to 2019, we repeatedly sampled the oligotrophic northwest Sargasso Sea in the vicinity of the Bermuda Atlantic Time-series Study site (BATS) to quantitatively follow known compounds within the pool of dissolved organic matter in the upper 1000 meters of the water column. Dissolved metabolite concentrations revealed patterns with depth and time with most metabolites showing surface enrichment and lower concentrations with increasing depth. Select metabolites displayed seasonal patterns throughout the year, and this seasonality was repeated in each of the years sampled. Concentrations of vitamins, including pantothenic acid (vitamin B5) and riboflavin (vitamin B2), increased annually during winter periods when mixed layer depths were deepest. During diel sampling, light-sensitive riboflavin decreased significantly during daylight hours. Metabolite concentrations over time at BATS showed less variability compared to a previous sample set collected over a broad latitudinal range in the western Atlantic Ocean. The metabolites examined in this study are all components of central carbon metabolism. By examining these metabolites at finer resolution and in a relatively long time series, we have insights into microbial biogeochemical activity, data which are fundamental to understanding the chemical response of marine systems to future changes in climate.

biochemistry↗

bioRxiv2022

Compartmentalization of specialized metabolites across vegetative and reproductive tissues in two sympatric Psychotria (Rubiaceae)

Premise of the studyThe specialized metabolites of plants are recognized as key chemical traits in mediating the ecology and evolution of sundry plant-biotic interactions, from pollination to seed predation. Intra- and interspecific patterns of specialized metabolite diversity have been studied extensively in leaves, but the diverse biotic interactions which contribute to specialized metabolite diversity encompass all plant organs. Focusing on two species of Psychotria shrubs, we investigate and compare patterns of specialized metabolite diversity in leaves and fruit with respect to each organs diversity of biotic interactions. MethodsTo evaluate associations between biotic interaction diversity and specialized metabolite diversity, we combine UPLC-MS metabolomic analysis of foliar and fruit specialized metabolites with existing surveys of leaf- and fruit-centered biotic interactions. We compare patterns of specialized metabolite richness and variance among vegetative and reproductive tissues, among plants, and between species. Key resultsIn our study system, leaves are involved in a greater number of host-specific biotic interactions than fruit, while fruit-centric interactions are more ecologically diverse. This was reflected in specialized metabolite richness - leaves contained more than fruit, while each contained over 200 organ-specific specialized metabolites. Within each species, leaf- and fruit specialized metabolite composition varied independently of one another across plants. Specialized metabolite composition exhibited stronger contrasts between organs than between species. ConclusionsAs ecologically disparate plant organs with organ-specific specialized metabolite traits, leaves and fruit can each contribute to the tremendous overall diversity of plant specialized metabolites.

plant biology↗

bioRxiv2022

Carbon content, carbon fixation yield and dissolved organic carbon release from diverse marine nitrifiers

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 less well known but may represent an as-yet unaccounted source of dissolved organic carbon (DOC) available to heterotrophic marine food webs. Here, we provide measurements of cellular carbon and nitrogen quotas, DIC fixation yields and DOC release of ten phylogenetically diverse marine nitrifiers grown in multiple culture conditions. All investigated strains released DOC during growth, making up on average 5-15% of the fixed DIC. Neither substrate concentration nor temperature affected 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 CO2 fixation, and due to lower observed yields in artificial compared to natural seawater medium. The results of this study provide 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.

ecology↗

bioRxiv2019

Pangenomics reveal diversification of enzyme families and niche specialization in globally abundant SAR202 bacteria

It has been hypothesized that abundant heterotrophic ocean bacterioplankton in the SAR202 clade of the phylum Chloroflexi evolved specialized metabolism for the oxidation of organic compounds that are resistant to microbial degradation via common metabolic pathways. Expansions of paralogous enzymes were reported and implicated in hypothetical metabolism involving monooxygenase and dioxygenase enzymes. In the metabolic schemes proposed, the paralogs serve the purpose of diversifying the range of organic molecules that cells can utilize. To further explore this question, we reconstructed SAR202 single amplified genomes and metagenome-assembled genomes from locations around the world, including the deepest ocean trenches. In analyses of 122 SAR202 genomes that included six subclades spanning SAR202 diversity, we observed additional evidence of paralog expansions that correlated with evolutionary history, and further evidence of metabolic specialization. Consistent with previous reports, families of flavin-dependent monooxygenases were observed mainly in the Group III SAR202, in the proposed class Monstramaria and expansions of dioxygenase enzymes were prevalent in Group IV. We found that Group I SAR202 encode expansions of racemases in the enolase superfamily, which we propose evolved for the degradation of compounds that resist biological oxidation because of chiral complexity. Supporting the conclusion that the paralog expansions indicate metabolic specialization, fragment recruitment and fluorescence in situ hybridization with phylogenetic probes showed that SAR202 subclades are indigenous to different ocean depths and geographical regions. Surprisingly, some of the subclades were abundant in surface waters and contained rhodopsin genes, altering our understanding of the ecological role of SAR202 in stratified water columns.\n\nImportanceThe oceans contain an estimated 662 Pg C of dissolved organic carbon (DOC). Information about microbial interactions with this vast resource is limited, despite broad recognition that DOM turnover has a major impact on the global carbon cycle. To explain patterns in the genomes of marine bacteria we propose hypothetical metabolic pathways for the oxidation of organic molecules that are resistant to oxidation via common pathways. The hypothetical schemes we propose suggest new metabolism and classes of compounds that could be important for understanding of the distribution of organic carbon throughout the biosphere. These genome-based schemes will remain hypothetical until evidence from experimental cell biology can be gathered to test them, but until then they provide a perspective that directs our attention to the biochemistry of resistant DOM metabolism. Our findings also fundamentally change our understanding of the ecology of SAR202, showing that metabolically diverse variants of these cells occupy niches spanning all depths, and are not relegated to the dark ocean.

genomics↗