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Limnology and Oceanography

Wiley

Preprints posted in the last 90 days, ranked by how well they match Limnology and Oceanography's content profile, based on 32 papers previously published here. The average preprint has a 0.03% match score for this journal, so anything above that is already an above-average fit.

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Carbon concentrating mechanism and growth response of the diatom P. tricornutum to changes in Zn and carbonate chemistry

Burdige, E.; Saito, M.; Hayden, M.; McIlvin, M.; Subhas, A.

2026-05-23 biochemistry 10.64898/2026.05.21.726966 medRxiv
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The efficiency of marine diatom carbon concentrating mechanisms (CCMs) play a critical role in photosynthesis and enable cells to maintain rapid growth rates under a variety of environmental conditions. To assess the plasticity of the model diatom P. tricornutums CCMs, specifically carbonic anhydrase (CA) enzymes and bicarbonate transporters, we measured growth response, bulk CA activity, and corresponding shifts in the proteome under a range of Zn and pCO2 conditions in culture. CA activity increased with Zn availability and decreased with pCO2. A positive growth effect was observed due to Zn addition and increasing pCO2 from 200 to 400 ppm, however growth rate decreased as pCO2 further increased to 750 ppm. Across the six treatments, the protein abundance of ISIP2A, which functions to bring Fe into the cell via a FeCO3 complex and is used as a biomarker for Fe stress, demonstrated an inverse relationship with [CO32-], consistent with its role as a phytotransferrin. Under conditions of Zn limitation ([Zn2+] = 0.3 pM), the cell appeared to allocate this metal away from CA, instead relying on a Mn-CA with a 100-fold lower intrinsic activity than that of the primary Zn-CA, as calculated using paired abundance-activity measurements. We further observed a continued increase in bicarbonate transport protein abundance after CA activity plateaued at 1.2x10-6 (reactions sec- 1cell-1), suggesting any deficit in DIC required to maintain high growth rates is accomplished through HCO3- uptake. We hypothesize that bicarbonate uptake and CO2 diffusion operate in tandem via CA enzymatic activity to supply adequate CO2 for photosynthesis.

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Differences between suspended and sinking particles regulate carbon flux in the upper mesopelagic during a Phaeocystis Bloom

Cisternas-Novoa, C.; Romanelli, E.; Passow, U.

2026-07-08 systems biology 10.64898/2026.06.09.731151 medRxiv
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Despite decades of research, the factors determining the sinking velocity of marine biogenic particles remain poorly constrained, and growing evidence suggests that particle composition and morphology are as important as size in determining particle fate. We compared characteristics of suspended and sinking particles at three depths below the mixed layer and within the layer of maximal flux attenuation during the decline of a Phaeocystis pouchetii bloom in the Labrador Sea using marine snow catchers. Biochemical and morphological characteristics of suspended and sinking particles always differed, with differences depending primarily on bloom stage, and depth accounting for comparatively less variation. Exopolymer particles played a key role, with the relative concentrations of transparent exopolymer particles consistently higher in the suspended than in the sinking particle fraction. In contrast, the partitioning of coomassie-stainable particles changed with the bloom stage, as a function of the Phaeocystis life cycle. Ballast minerals played a negligible role during the late-bloom and bloom-decline stages, and their relative importance increased during the non-bloom stage. The C:N ratio was lower in suspended than sinking particles, with differences in morphological measures depending on bloom stage. Our findings emphasize that export potential is driven not only by particle size, but also by bloom stage, which is closely linked to plankton community composition and plays a key role in the timing and magnitude of carbon flux in the upper mesopelagic. Further, this work highlights the important and diverse roles of exopolymers in regulating carbon flux.

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Glacier-induced upwelling shapes microbial communities in Arctic marine systems

Spence, J. S.; Bertrand, E. M.; White, P. L.; Parrott, C. M.; Waterman, S.; Didier, D.; Roberts, M. E.; Hamilton, A. K.; Cavaco, M.; Noah, T.; Mahmoudi, N.; Konhauser, K.; Bhatia, M. P.

2026-05-13 microbiology 10.64898/2026.05.12.724575 medRxiv
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The Canadian Arctic Archipelago (CAA) is warming at an unprecedented rate, leading to sea ice loss and glacial retreat. Marine-terminating (tidewater) glaciers can fuel summertime marine productivity by delivering nutrient-rich deep waters via upwelling to the surface ocean. While the impact of glacier-induced upwelling has been well-studied in the context of phytoplankton and primary productivity, its effects on broader marine microbial communities remain poorly understood. We investigated how glacier-driven upwelling shapes marine microbial (bacterial and archaeal) communities across a series of sites in the CAA. At upwelling sites, the upper 50 m of the water column exhibited elevated nutrient concentrations and physical characteristics that resembled deeper waters, which were associated with differences in microbial community composition relative to non-upwelling sites. Our results indicate that upwelling influences microbial communities in surface waters in two ways. It directly introduces typically deeper-water-associated taxa into surface waters and reshapes ecological niches by enhancing nutrient supply and stimulating primary production, indirectly driving changes in microbial communities. The enrichment of Candidatus Nitrosopumilus, a deep water nitrifier, likely affects nitrogen cycling and raises the possibility of active nitrification in surface waters. Likewise, the increased abundance of taxa known to be associated with phytoplankton-derived organic matter in upwelling regions suggests an enhanced capacity to process organic matter generated from elevated primary productivity. Ultimately, as tidewater glaciers continue to retreat, the resulting changes in the glacially-driven upwelling regime will likely shift marine microbial communities towards assemblages adapted to less productive ecosystems, with implications for nutrient cycling in these systems. ImportanceClimate change has a disproportionate impact on the Arctic, with rising temperatures causing increased marine-terminating glacier retreat and changes in the marine water column structure. The consequent loss of the ability of these glaciers to upwell deep water to the surface ocean results in a reduction of nutrient delivery and mixing in these ecosystems. Previous work has highlighted the importance of marine-terminating glaciers in sustaining phytoplankton productivity during the summer season through this delivery of deep-water nutrients to the surface ocean. The impact of glacially-induced upwelling on marine bacterial and archaeal communities, however, remains underexplored. We found that in regions with glacially-driven upwelling, the surface ocean showed enrichment of phytoplankton-associated taxa and nitrifiers commonly associated with deep waters. This work underscores the role of glacially-driven upwelling in structuring both microbial communities and nutrient cycling, suggesting that glacier loss could reshape community composition and biogeochemical processes in a rapidly changing Arctic.

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Virus-grazer interplay enhances virus production, particle aggregation, and trophic efficiency during infection of Prochlorococcus

Lindell, D.; Carlson, M. C.; Weissenbach, J.; Kirzner, S.; Sulcius, S.; Hulata, Y.; Sabehi, G.; Goldin, S.; Sacks, J. S.; Bjorkman, K. M.; Dugenne, M.; Zborowsky, S.; Linney, M. D.; Beckett, S. J.; Tahan, R.; Demory, D.; White, A. E.; Weitz, J. S.; Karl, D. M. M.; Armbrust, V.; Ingalls, A. E.; Ribalet, F.; Caron, D. A.

2026-06-11 microbiology 10.64898/2026.06.10.731369 medRxiv
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Viruses and grazers are fundamental agents of mortality in the oceans, impacting phytoplankton populations and organic matter cycling. Although viruses and grazers co-occur in nature, they are typically studied in isolation in laboratory experiments, limiting our understanding of their combined ecosystem impacts. Here, using a simplified ecosystem approach, we investigated individual and combined effects of the T7-like cyanopodovirus, P-SSP7, and the protistan grazer, Paraphysomonas bandaiensis, on the abundant marine cyanobacterium, Prochlorococcus MED4, and co-occurring non-photosynthetic heterotrophic bacteria (bacteria from here on). We observed that, individually, viruses and grazers caused substantial Prochlorococcus mortality. Viral lysis also triggered increases in damaged Prochlorococcus cells, dissolved organic matter release, and bacterial growth, while grazing reduced bacterial abundances. When grazers and viruses were combined, Prochlorococcus mortality was lower than expected from the sum of their individual effects. Contrary to expectations, this reduced Prochlorococcus mortality did not result in fewer viruses or grazers. Instead, virus-grazer-Prochlorococcus interplay resulted in greater virus production, maintenance of grazer growth, and a dramatic increase in particle aggregation. Our results reveal trophic cooperation and efficiency in which competition between viruses and grazers was likely mitigated, with virus progeny production enhanced by grazers, and grazer growth sustained through a shift to alternative food sources (bacteria, damaged cells, aggregates) secondarily derived from Prochlorococcus following viral lysis. The synergistic enhancement of particle aggregation via grazer-virus-phytoplankton interplay observed with the small buoyant Prochlorococcus phytoplankter underscores the importance of food web interactions for the flow of phytoplankton-fixed carbon within, and export from, the photic zone. SignificanceViruses and grazers both use phytoplankton as a resource for reproduction. In a simplified experimental system with Prochlorococcus, an important primary producer in the oceans, we found that the interplay between viruses and grazers led to reduced mortality of Prochlorococcus. Despite this reduced mortality, virus-grazer interactions resulted in elevated virus production and a dramatic increase in organic matter aggregation. Furthermore, grazer abundance was not affected by this interplay, likely due to the transfer of organic matter from Prochlorococcus to bacteria and aggregates, which the grazers could consume as alternative food sources. These findings provide insights into the complexity of ecosystem interactions and how they impact the fate of organic matter fixed by phytoplankton in the oceans.

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Shifts in consumer assemblages are linked to nutrient availability and ecosystem metabolism

Sadler, I.; Stanley, A.; Narr, C. F.

2026-04-27 ecology 10.64898/2026.04.23.720454 medRxiv
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Nutrient availability, ecosystem productivity, and consumer assemblages are intricately linked through complex interactions and feedbacks. Nutrients influence the diversity and functional roles of consumers via shifts in resource quality and quantity, and consumers can alter ecosystem production and nutrient availability. However, our understanding of how characteristics of consumers respond to and influence concomitant shifts in nutrient availability and production is limited. We quantified the response of well-studied consumer assemblages (benthic invertebrates and zooplankton) to realistic nutrient loads that altered gross primary production (GPP) and ecosystem respiration (ER). We fertilized 14 outdoor experimental ponds for 2 months and monitored total water column carbon (TC), nitrogen (TN), and phosphorus (TP), GPP, ER, and net ecosystem production (NEP) weekly. Then, we evaluated how fertilization and the variation in nutrients and metabolism caused by fertilization were related to shifts in consumer assemblages. Fertilization increased water column TN and TP and reduced TC:TP ratios, TN:TP ratios, and rates of GPP and ER. However, consumer assemblages were more tightly linked to variation in nutrient availability and production across ponds than to fertilization. Greater declines in benthic diversity occurred in ponds with higher average TN:TP ratios during the experiment. Consistent with predicted effects of cladocerans on nutrient availability, shifts in cladoceran abundances were positively associated with average water column TN:TP ratios during the experiment. Finally, elevated GPP and ER were associated with greater increases in the abundance of benthic invertebrate predators, suggesting the possibility of top-down control. Our study highlights the critical role of consumer-mediated processes in the interaction between nutrient availability and production. Manuscript HighlightsO_LIFertilization reduced pond gross primary production and ecosystem respiration rates. C_LIO_LIInvertebrate predator abundance was inversely related to gross primary production. C_LIO_LIShifts in consumer assemblages were tightly linked to nutrients and production. C_LI

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The effects of thermal alteration on organic matter bioavailability in deeply buried marine sediments

McNichol, S. M.; Shah Walter, S. R.; Teske, A. P.; Mahmoudi, N.

2026-06-17 microbiology 10.64898/2026.06.16.732717 medRxiv
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A substantial fraction of marine sediments experience elevated temperatures due to burial or hydrothermal activity. These conditions can fundamentally reshape both microbial activity and the chemical nature of sedimentary organic matter (OM). Laboratory incubations have demonstrated that moderate heating of marine sediments can lead to the production of labile organic compounds such as acetate, however, it remains unclear whether heating alters the bioavailability of the remaining OM pool. In this study, we experimentally tested the effect of temperature on the bioavailability of OM through a series of bioreactor experiments using deeply buried sediment collected from Guaymas Basin (Gulf of California). We measured acetate concentrations in sterilized Guaymas Basin sediments before and after artificial heating (70{degrees}C for 7 days) to quantify abiotic acetate generation. We then conducted incubations of a model marine bacterium with sterilized, artificially heated sediment and tracked respired CO2 production and its associated 13C and 14C signatures. Our study revealed that sediment depth and hydrothermal history strongly control abiotic acetate production, with higher acetate yields from shallower, cooler sediments. Respiration rates in control and heated sediment incubations were nearly identical, indicating that heating does not measurably alter the bioavailability of bulk sedimentary OM. Moreover, the {delta}13C values of respired CO2 were indistinguishable between control and heated sediment incubations while the {Delta}14C values were more depleted in the first 24 hours in incubations with heated sediment. This transient offset suggests that low-temperature heating mobilizes a small pool of older material due to desorption of mineral-bound OM without altering overall bioavailability. Our findings shed light on the role of thermal alteration in shaping carbon cycling in marine sediments by influencing how OM is made available to sedimentary microorganisms.

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Lifestyle modulates riverbed diazotrophy: from abundance and activity to diversity of free-living and biofilm-associated N2 fixers

Lander, R.; Villamarin, C.; Siebner, H.; Arnon, S.; Bar Zeev, E.

2026-04-29 microbiology 10.64898/2026.04.27.721245 medRxiv
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Despite the critical role that biological N2 fixation plays in controlling primary and secondary production in aquatic ecosystems, freshwater subsurface diazotrophy has received minimal attention. Here, we quantified N2 fixation rates, diazotroph abundance, and diversity in the hyporheic zone across seasons and redox regimes, distinguishing between free-living and biofilm-associated lifestyles. Samples were collected from aerobic and suboxic strata of the Jordan River streambed and incubated in microcosms for 24 h with dissolved 15N2 under ambient conditions. Immunolabeling the nitrogenase enzyme coupled with flow-cytometry revealed that diazotrophs accounted for [≤]9.6 % of total bacterial abundance but were consistently enriched in biofilms. These biofilms supported 377-fold higher N2 fixation rates per-cell than free-living diazotrophs. Confocal laser scanning microscopy captured thicker (several tens of microns) complexes of extracellular polymeric substances (EPS) encasing aerobic biofilms than suboxic, especially during winter. Biofilm-associated communities retained high abundances ([≤]21 %) of facultative and obligate anaerobes in oxic zones. Compared to those in biofilms, the abundance, N2 fixation rates, and diversity of the free-living fraction varied much more across both aerobic and suboxic zones. nifH sequencing unveiled the dominance of Pseudomonadota and Thermodesulfobacteriota across all communities. Overall N2 fixing activity, when converted to volumetric units, was 1.5 x 105-fold greater in the streambed than the overlying water. These findings identify the hyporheic zone as an active hotspot for benthic diazotrophy and intimate the centrality of microbial lifestyle in determining how diazotrophs persist and remain highly active in fluctuating redox and nutrient conditions in the freshwater ecospace.

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Lake ecosystem responses to runoff variability across time and space

Langenheder, S.; Mesman, J. P.; Kreuter, N.; Kothawala, D.; Agreda-Lopez, G.; Ari, A.; Berger, S. A.; Bernal, S.; Buttyan, B.; Bick, B.; Carabal, N.; Catalan, N.; Charmpila, E. A.; Colom Montero, W.; Erturk Ari, P.; Elfferich, I.; Exner, J.; Gergacz, B.; Gray, E.; Happe, A.; Jiao, C.; Jones, K.; Karakaya, N.; Kulas, A.; Lupon, A.; Mangold, C.; Mendoza-Lera, C.; Nejstgaard, J. C.; Oppong, J.; Pedregal-Montes, A.; Perujo, N.; Rankinen, J.; Rutting, T.; Sjostedt, J.; Striebel, M.; Symiakaki, K.; van Dam, E.; Wentritt, S.; Yaqoob, M. M.; Yildiz, K.; Sassenhagen, I.

2026-06-19 ecology 10.64898/2026.06.18.729811 medRxiv
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Inland waters in the Northern Hemisphere are experiencing increased annual runoff due to higher overall precipitation as well as intensified short-term events such as heavy rainfall, floods and storms. These events affect the total loading and variability of inputs of allochthonous, coloured dissolved organic matter (cDOM) and inorganic nutrients into lakes. Previous studies have shown that increased total cDOM and inorganic nutrient loads affect phytoplankton biomass and metabolic rates, but it is unknown how the effects of different cDOM and nutrient pulse scenarios are modified by spatial and seasonal differences in lake characteristics. Here, we conducted a coordinated, standardized mesocosm experiment across three lakes with different ambient cDOM and nutrient concentrations. In two of these lakes, the experiment was implemented in two seasons. The same total amounts of cDOM, nitrate and phosphate were added to all mesocosms, but in pulses that differed in intensity and frequency. We found that pulse intensity and frequency affected chlorophyll a and phycocyanin concentrations and metabolic rates, i.e. gross primary production and respiration, differently. Specifically, more pronounced effects were found in response to the extreme pulse scenario compared to those with more frequent, smaller pulse additions. Furthermore, the effects were mainly temporary and varied more among lakes than between seasons. The clearest differences between the extreme and more gradual runoff scenarios were found in the lake with the lowest background cDOM and nitrate concentrations, likely because lower light limitation and possibly stronger initial N-limitation caused a faster response to the nutrient addition. Our results highlight that both antecedent lake conditions and characteristics of runoff events can affect phytoplankton biomass and metabolic rates and that comparative experimental approaches are needed to reveal the complexity of the responses.

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A low-cost method for collecting and measuring sinking speed of particles in coastal zones

Lemke, J.; Spilling, K.

2026-06-15 biochemistry 10.64898/2026.06.11.730824 medRxiv
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Sinking marine particles is a key process regulating carbon export through the biological carbon pump, yet direct measurements of sinking dynamics remain limited in many coastal environments. One barrier is that most existing approaches require expensive instrumentation and large research platforms. Here, we present a low-cost, modular method for concentrating fast-sinking particles and measuring their individual sinking velocities under controlled conditions. This combines large settling tanks (110 L) for field-based particle fractionation with a video-based tracking system that quantifies the sinking behavior of natural marine particles. The particle sinking speed chamber is surrounded on three sides by a temperature-controlled water chamber, minimizing the problem of advection during measurements. The post-processing Python script delivers sinking velocity, particle size, circularity, and RGB-based properties for large numbers of particles. The method accuracy was validated using reference beads with known theoretical sinking velocities derived from Stokes law. Field deployments in the Baltic Sea demonstrated successful enrichment of fast-sinking particles and stable operation from both a research vessel and a small boat. Compared to existing methods, the approach substantially reduces logistical and financial barriers while maintaining particle-resolved measurements and compatibility with complementary biogeochemical analyses. This enables a broader observational coverage of sinking particle processes across environments that are currently underrepresented in carbon export studies.

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An ice-bucket challenge: investigating ice algae physiology in laboratory microcosms.

Baker, M. L.; Forss, E.; Kolzenburg, R.; Collins, S.; Kranz, S. A.

2026-07-13 ecology 10.64898/2026.07.10.737583 medRxiv
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John Raven pioneered the field of algae ecophysiology, advancing our understanding of cellular resource economics, carbon acquisition, and energy allocation. His work laid the foundation for investigating integrative physiology, linking growth-survival trade-offs across diverse environments. The sea ice habitat provides an excellent framework to continue the research John championed. With steep temperature-salinity gradients, algae survival requires a shift in physiology that we are only beginning to understand. We developed two small scale, reproducible icecosms to investigate physiological changes associated with incorporation into sea ice and survival potential post-melt. Fragilariopsis cylindrus and Nitzschia frigida, known for their association with the ice environment, and Porosira glacialis, known for its association with the ice edge, were used to mechanistically link physical properties with algal physiology and post-melt survival. We observe incorporation into the ice of F. cylindrus and N. frigida alongside vertical photophysiological profiles of F. cylindrus revealing inhospitable conditions in the top compared to the bottom layers of ice. N. frigida and P. glacialis remain viable within the ice and retain the capacity to seed populations following melt. Our results establish icecosms as experimental framework to investigate ecophysiological responses of sea ice algae and provide a foundation toward ecological and evolutionary questions.

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Rapid shallow-water saturation and deep-water expansion of an invasive freshwater ecosystem engineer in a deep European lake

Hofstetter, L.; Mueller, T. M.; Bourqui, M.; Burlakova, L. E.; Cristante, Z. C.; Karatayev, A. Y.; Kessler, S.; Narwani, A.; Santos, J. L.; Sturm, L.; Wellauer, N.; Spaak, P.; Weber, A. A.-T.

2026-06-27 ecology 10.64898/2026.06.26.734794 medRxiv
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Quagga mussels (Dreissena rostriformis bugensis) are ecosystem engineers that can alter nutrient cycling, benthic-pelagic coupling, and food-web structure in deep lakes. Although their invasion trajectories are well documented in the Laurentian Great Lakes in North America, depth-specific population dynamics remain poorly resolved in recently invaded European perialpine lakes. We analyzed five annual lake-wide surveys (2021-2025) from 54 stations spanning 2.4-253 m depth in Lake Constance to quantify changes in quagga mussel density, biomass, and shell-length distribution. Contrary to expectations of lake-wide exponential growth, shallow-water populations (< 20 m) showed no significant increase during the study period and appear to have reached carrying capacity before monitoring began. In contrast, densities increased monotonically at intermediate depths (40-125 m), indicating ongoing expansion into deeper strata. Mean shell length declined with depth, and size distributions in shallow waters shifted toward larger individuals, consistent with a transition from active recruitment to somatic growth of established mussels. Compared with the Laurentian Great Lakes, Lake Constance already has substantially higher shallow-water biomass, whereas deeper invasion trajectories are broadly similar. These results show that quagga mussel invasion in deep European lakes can combine rapid littoral saturation with slower profundal expansion, complicating direct transfer of predictions from the Great Lakes. Continued depth-stratified monitoring will be essential for anticipating future ecosystem effects in perialpine lakes.

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Environmental microbial communities and host selection shape larval microbiomes

Hendricks, S. F.; Tan, A. L.; Williams, A. G.; Buckley, K. M.; Strader, M. E.

2026-05-15 ecology 10.64898/2026.05.14.725214 medRxiv
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Ocean warming is altering abiotic environments and biotic interactions experienced by marine organisms, where sensitive early developmental windows occur in biologically complex seawater communities. The impact of these interactions on developmental processes and fitness in hosts is not well understood, but likely contingent on the establishment of a host-associated microbiome. Here, we hypothesize that temperature and microbial exposure during embryogenesis influence larval microbiome assembly and host morphology. Strongylocentrotus purpuratus embryos were raised in low microbial richness (LMR) or high microbial richness (HMR) seawater at ambient (14 {degrees}C) or elevated (18 {degrees}C) temperature, then collected at 2, 4, and 6 days post-fertilization (dpf) following multiple feedings. Higher microbial diversity was observed in larvae that developed in HMR seawater when compared to LMR. Differences in relative abundances of dominant microbial families between seawater and larvae suggest some degree of host selectivity in microbiome assembly. Temperature did not strongly alter microbiome composition, but both temperature and microbial condition led to differences in larval morphology by 6 dpf, potentially due to enrichment of microbes with chemoheterotrophic functions. By linking how temperature and microbial communities interact with host development, we contribute novel insights into how early-life environmental conditions impact holobiont formation and morphology. One sentence summaryEarly developmental temperature and microbial conditions shape larval microbiome establishment and morphology.

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Short term effects of salinization on the plankton community of an oligotrophic mountain lake

Ptacnik, R.; SalInvade group, lead by Izabele Suikate, ; PP-TOX group, lead by Elisabeth Varga,

2026-07-13 ecology 10.64898/2026.07.10.737327 medRxiv
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Freshwater salinization is of increasing concern for integrity and functioning of freshwater habitats worldwide. Experiments so far often have studied drastic salt additions, while gradient designs have been performed less commonly. We tested the effect of freshwater salinization in a mesocosm exposing the plankton community of the oligotrophic Lake Lunz, Austria, to a four-fold salinization gradient (control, 0.2, 1, a 5 ppt salt). Salinity was manipulated in a factorial design with enrichment, with 10 g L-1 and 30 g L-1 phosphorus, resulting in 8 treatments with 3 replicates each. We followed the effects of salinization on diversity, community composition and resource use over 36 days. Community composition was assessed by amplicon sequencing, Diversity loss and community turnover followed upon salt addition. All levels of salinization caused pronounced changes in community composition, with 5 ppt causing the most drastic changes. Salinization caused trophic downgrading by kicking out especially protistan consumers and rotifers, while some green algae and chrysophytes were especially tolerant, resulting in reduced phylogenetic and functional diversity with increasing salinization. In line with reduced top down control, salinization affected temporal variability in chlorophyll-a (chl-a) and resource use (RUE), with higher salinity causing more extreme fluctuations in chl-a and RUE. Enrichment overall aggravated salinization, enhancing temporal turnover and temporal fluctuations in resource use.

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Microbial Growth in an Enceladus Ocean Analog Medium Informed by Mineral Stability Modeling

Elkassas, S. M.; Ely, T.; Zhivkova, T.; Patterson, A.; Weeks, K.; Mitchell, S.; Hayes-Guastella, L.; Nathan, V.; Serres, M.; Shock, E.; Girguis, P.; German, C.; Klein, F.; Seewald, J.; Huber, J. A.

2026-07-01 microbiology 10.64898/2026.06.29.735333 medRxiv
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Evidence from the Cassini mission confirmed that Saturn's moon Enceladus hosts a subsurface alkaline ocean where rock-water reactions may generate redox disequilibria capable of supporting microbial metabolisms. To investigate potential microbial survival under simulated Enceladus ocean conditions, we used thermodynamic modeling to develop a salt formulation consistent with one possible Enceladus ocean composition and supplemented it with putative microbial energy sources to create a growth medium. The medium was inoculated with samples from diverse ocean world analog environments on Earth to determine which microorganisms could persist under Enceladus-like conditions. The microorganisms persisting in this geochemically bounded medium were heterotrophic, metabolically versatile bacteria with low carbon requirements. Genomic and physiological analyses further showed the presence of multiple stress-response pathways, sodium- based bioenergetic systems, osmoregulation strategies, and other adaptations consistent with survival in alkaline, low-nutrient settings. These results suggest that some stress-tolerant heterotrophic bacteria may serve as useful model organisms for life in Enceladus' subsurface ocean. These findings demonstrate the value of geochemically modeled media as a framework for constraining habitability, identifying relevant biosignatures, and probing potential microbial survival strategies beyond Earth.

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Seasonal microbiome community dynamics in the massive coral Porites lobata impacted by sedimentation

Berg, J. T.; Fifer, J.; Davies, S.; Bentlage, B.

2026-05-12 ecology 10.64898/2026.05.08.723747 medRxiv
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Near-shore coral reefs in southern Guam (Mariana Islands) experience severe sedimentation, in particular during the wet season when rainfall and erosion are high. We sampled fragments of the reef-forming coral Porites lobata from opposite ends of a sedimentation gradient in Fouha Bay, southern Guam, during dry and wet seasons. Using DNA metabarcoding, we characterized the diversity and composition of P. lobata-associated Symbiodiniaceae and bacterial microbiome communities. As in many species of Porites, Symbiodiniaceae communities of P. lobata were dominated by variants of Cladocopium C15 with sites showing differences in Symbiodiniaceae communities attributable to variation in these Cladocopium C15 variants. Bacterial microbiomes of P. lobata were dominated by Endozoicomonadaceae, a family of putative coral bacterial endosymbionts involved in nutrient cycling. Site and seasonal differences in bacterial diversity and community composition were apparent. In close proximity to the mouth of the river draining into Fouha Bay, bacterial diversity was highest during the wet season when sedimentation is generally severe. Microbiome reorganization in response to sedimentation may explain this result, but we also found overrepresentation of bacteria associated with terrestrial origin close to the river mouth and/or during the wet season. Together these patterns highlight that coral Symbiodiniaceae and bacterial communities are both spatially and temporally structured in this disturbed system. IMPORTANCEThis study provides a time series dataset of coral-associated microorganisms, including dinoflagellate algae and bacteria, from a tropical bay impacted by sedimentation that results from upstream erosion of disturbed soils. Characterizing temporal patterns of coral-associated microbes provides insights into the dynamic nature of these communities. While microbiome variability across sites and seasons may be a result of acclimatization to different environmental conditions, we identified bacterial groups of putative terrestrial origin in sampled coral microbiomes that may have been exported from eroded soils to the near-shore reef. Considering that disturbed soils act as hotspots for the proliferation of potentially harmful substances, such as antimicrobial resistance genes, understanding microbial community connections at the marine-freshwater-terrestrial interface is an important step toward evaluating environmental impacts across connected ecosystems from ridge to reef.

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A closer look at plankton: potential interactions inferred from centimeter-scale in situ observations

Panaiotis, T.; Irisson, J.-O.; Freilich, M.; Cael, B. B.

2026-05-20 ecology 10.64898/2026.05.18.725820 medRxiv
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Plankton are essential to marine ecosystems, supporting food webs and mediating biogeochemical processes such as carbon export to depth. Their spatial distribution influences ecosystem dynamics and serves as an indicator of environmental change. Although drifting plankton could be expected to exhibit random distribution, numerous studies have revealed significant heterogeneity in their spatial patterns. However, very few studies targeted plankton distribution at the centimeter scale in situ, despite its importance for understanding biological processes. We argue that centimeter-scale distances in plankton could reveal potential ecological interactions. Using an extensive in situ dataset of 18 million planktonic organisms collected by the In Situ Ichthyoplankton Imaging System (ISIIS), which images multiple organisms simultaneously and preserves their positions in the water column, we analyzed centimeter-scale distances in plankton. By comparing observed distances with those expected under a random distribution, we assessed potential interactions at three levels: among all organisms, within plankton groups and across groups. Our results show that planktonic organisms exhibit non-random distributions at the centimeter scale, with smaller distances than expected, suggesting potential ecological interactions. Notably, distances up to 11 cm were the most informative, which is much larger than typical interaction distances in plankton. Additionally, observed distances were compatible with a simple attraction model. Finally, we propose the non-randomness of distances as a novel metric of interaction strength in plankton ecological networks and compare it against classical empirical or co-occurrence networks. These results offer new insights into in situ interactions and how they shape plankton distribution at centimeter scale. Significance statementThis study reveals that planktonic organisms exhibit non-random spatial distributions at the centimeter scale, highlighting the importance of ecological interactions in shaping their distribution at this scale. By analyzing an extensive in situ plankton imaging dataset, we introduce a novel metric of interaction strength based on the non-randomness of distances between organisms, and compare it to common interaction metrics. These findings challenge the traditional view of plankton as passive drifters by highlighting that their distribution at microscale is shaped not just by physical processes such as turbulence but also by ecological interactions. Author contributionsJOI contributed to data acquisition. TP processed the data under the supervision of JOI. TP, JOI and BBC designed the study. TP conducted the analyses under the supervision of MF, JOI and BBC. TP wrote the initial draft of the manuscript. All authors contributed to the interpretation of results, supported manuscript preparation and approved the final submitted version.

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Seasonal patterns in Synechococcus pigment diversity at two temperate sites with contrasting oceanic regimes

Dufour, L.; Faure, E.; Partensky, F.; Mattei, F.; Uitz, J.; Petit, F.; Vellucci, V.; Golbol, M.; Ratin, M.; Gouriou, B.; Gachenot, M.; Clairet, J.; Farrant, G. K.; Hoebeke, M.; Corre, E.; Antoine, D.; Baudoux, A.-C.; Bigeard, E.; Bureau, S.; Castel, J.; Chambouvet, A.; Couet, D.; Cre hriou, R.; de Vargas, C.; Dimier, C.; Le Gall, F.; Guillou, L.; Henry, N.; Rigaut-Jalabert, F.; Jeanthon, C.; Romac, S.; Simon, N.; Szymczak, J.; Trellu, C.; Walde, M.; Hickman, A.; Dutkiewicz, S.; Kehoe, D. M.; Not, F.; Thiebaut, E.; Garczarek, L.

2026-07-13 microbiology 10.64898/2026.07.10.737521 medRxiv
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Competition for light has driven extensive pigment diversification among phytoplankton species, yet how this diversity shapes their spatiotemporal distribution in the field has been little studied so far. The cyanobacterium Synechococcus is an ideal model for addressing this issue, since this group has colonized most light spectral niches in marine environments. Here, we used an approach based on marker read recruitment from metagenomes to analyze the seasonal succession of Synechococcus pigment types (PTs) at two time-series stations off French coasts exhibiting contrasting oceanic regimes. Marked seasonality was observed at both sites. The shallow, permanently mixed English Channel site SOMLIT-Astan was characterized by an alternation between green-light specialists (PT 3a) peaking in spring, and chromatic acclimaters type A (PT 3dA) accounting for most of the Synechococcus community in winter. In contrast, the pigment diversity was much higher at the deep Mediterranean station BOUSSOLE. In the upper layer, the two main PTs were the blue light specialists (PT 3c), which dominated the community in summer and fall, and PT 3dA cells, which were more abundant in spring. The third most abundant PT was chromatic acclimaters type B (PT 3dB), which accounted for up to 15% of the surface community in late fall. Strikingly, PT 3dA was dominant at depth during most of the year. Multivariate analyses between PT abundances, clade abundances and environmental factors, notably water color indexes, suggested new associations between PTs to specific clades and ecological niches. This study provides novel insights for refining distribution models of Synechococcus PTs and phytoplankton groups in general.

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The natural history of bacterial bloomers in a decade-long time series

Deulofeu Capo, O.; Garcia-Comas, C.; Rey-Velasco, X.; Auladell, A.; Logares, R.; Garces, E.; Ferrera, I.; Sanchez, O.; Gasol, J. M.; Sebastian, M.

2026-07-01 microbiology 10.64898/2026.07.01.735832 medRxiv
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Bacterial bloomers,populations that experience rapid and significant increases in abundance in response to environmental triggers, briefly dominate marine microbial communities, potentially impacting the ecosystem by channeling large amounts of nutrients and affecting carbon fluxes. Due to their ephemeral nature, bacterial bloomers are challenging to capture, and it remains unknown whether they are restricted to specific taxonomic groups or whether they exhibit recurrent patterns. We analyzed a decade-long time series from the Blanes Bay Microbial Observatory (BBMO, NW Mediterranean Sea) to investigate bacterial bloomers in two size fractions (free-living (0.2-3 um) and particle-attached (3-20 um) communities. We identified 57 Amplicon Sequence Variants (ASVs), less than 1% of the total bacterial richness, exhibiting recurrent or chaotic blooming-like behavior. Bloomers spanned diverse phyla, though some taxonomic coherence appeared within families containing multiple blooming taxa. Monthly sampling detected bloom events on average 4.6 +- 1.9 times per year across both size fractions. Once seasonality was accounted for, blooms showed weak associations with biological and physicochemical variables, likely a consequence of monthly sampling resolution. Nonetheless, a marked shift in the blooming community within the particle-attached size fraction coincided with ecosystem disturbances from the nearby harbour restoration, suggesting that bloomers may act as disturbance sentinels. Metagenomic data showed that blooms led to marked shifts in the community functional potential. Overall, our findings underscore the importance of investigating bloom dynamics to understand microbial contributions to biogeochemical cycles and stress the need for higher-frequency sampling to accurately capture these transient but ecologically relevant events.

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Renewable energy caves: water replenishment holes in offshore monopiles create novel marine habitats

Kornau, L. M.; Leutelt, B.; van Sluis, C. J.; Bruinsma, N.; Mascini, M. D.; Olie, R. A.; Jacobs, F. A.; van den Akker, S.; de Haan, E.; van Onselen, E.; Strigin, N.; Nijland, R.; Coolen, J. W.

2026-06-05 ecology 10.64898/2026.06.03.729839 medRxiv
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The expansion of offshore renewable energy introduces artificial hard substrate, but the ecological effects may change depending on design features. For example, water replenishment holes, implemented for internal water refreshment and corrosion control, also allow colonisation of previously inaccessible monopile interiors, creating a novel semi-enclosed habitat. This study compared epifaunal communities on the interior and exterior walls of four water replenishment hole-equipped monopiles in the southern North Sea and modelled internal water quality to better understand factors shaping these communities. Vertical video transects were used to quantify percentage cover along depth gradients, while a coupled hydrodynamic-water quality model predicted vertical patterns in dissolved oxygen and particulate organic carbon over one year. Monopile interiors function as semi-enclosed, cave-like habitats with distinct environmental conditions, including darkness, restricted water flow, and vertical gradients in dissolved oxygen and particulate organic carbon. Compared to the external cover, interior communities showed reduced dominance of typical North Sea hard-substrate taxa and increased heterogeneity, with higher contributions of sponges, calcareous tube worms, and brittle stars, resembling communities reported from marine cave environments. Overall epifaunal cover was lower on the interior walls and broadly reflected vertical patterns in water quality. Organic matter accumulated on the interior seafloor, with indications of microbial mat formation. These findings suggest that the internal environmental conditions influence community development. Water replenishment hole design may therefore shape community composition inside monopiles, with implications for possible use as nature-inclusive design and environmental management. Graphical abstract O_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=78 SRC="FIGDIR/small/729839v1_ufig1.gif" ALT="Figure 1"> View larger version (43K): org.highwire.dtl.DTLVardef@17eaed4org.highwire.dtl.DTLVardef@7e631org.highwire.dtl.DTLVardef@99b085org.highwire.dtl.DTLVardef@142376_HPS_FORMAT_FIGEXP M_FIG C_FIG

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Depth-dependent eDNA abundances across ecosystems inform deep-sea sampling strategies

Herrera, S.; Govindarajan, A. F.; Andruszkiewicz Allan, E.; Francolini, R.; Frates, E.; McCartin, L.; Pittoors, N. C.; Sengthep, M.; Stover, S.; Vohsen, S.; Yang, N.

2026-05-14 ecology 10.64898/2026.05.12.724363 medRxiv
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Environmental DNA (eDNA) surveys are increasingly used to assess marine biodiversity and inform deep-sea environmental decision-making, including mineral resource management and fisheries oversight. Yet standard low-volume protocols inherited from coastal work may be inadequate at depth, and no quantitative framework links depth and ecosystem context to defensible filtration volume targets. We compiled 841 eDNA samples from eight expeditions across the North Atlantic, Wider Caribbean, and Pacific (surface to 4000 m) to quantify how recoverable eDNA scales with depth and surface productivity, and to derive depth- and productivity-aware sampling targets. Total eDNA concentration declined with depth as a power law, with attenuation exponents (b) modulated by surface productivity: most gradual in eutrophic waters (b = 0.67), intermediate in mesotrophic (b = 0.90), and steepest in oligotrophic systems (b = 1.25); volume-weighted models explained 66-88% of the variance. At a fixed extract-concentration target, required filtration volumes diverged ~7-fold between oligotrophic and eutrophic systems at 200 m and ~38-fold at 4000 m. Conventional Niskin sampling, therefore, undersamples deep-sea biodiversity, particularly in mid- to low-productivity systems. Among laboratory parameters, the assay-specific extract-concentration target exerted greater leverage on required volume than extraction efficiency or elution volume. Volume-aware sampling paired with optimized recovery should be routine in deep-sea eDNA surveys.