Norman Göbeler’s research while affiliated with University of Helsinki and other places

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Publications (8)


Overview of Hotfloor system from, the (a) surface (drone picture by, Alf Norkko) and (b) underwater at the seafloor. The temperature control unit, consisting of 10 thermostats and the electric heater, is placed on shore ①. The electric heater is connected to a closed water circuit distributing warmed water through about 40 m long tubing ② to the weighted benthic chambers placed on the seafloor (b). About 10 m of tubing are coiled inside the chamber walls ③ to ensure a gentle heat exchange with the surrounding water. The chamber lid is equipped with various ports for sampling, a circulatory pump with diffuser ④ and sensors ⑤.
Overview of daily average temperature of induced marine heatwaves (MHW) treatment and control at ambient temperature (dashed line) and sampling occasions (numbers 1–3). The line colours show the MHW categories (yellow = moderate, orange = strong, red = severe) when exceeding reference values and the blue line shows the climatological mean based on period 1931–2020 (Goebeler et al., 2022). The thresholds for the MHW categories here are, for example on the 8 July 2021, for Category I ‘moderate’ 18.5°C, for Category II ‘strong’ 22.2°C, for Category III ‘severe’ 25.9°C. During 1—‘Acclimation’ all chambers (n = 10) were at ambient temperatures of about 18°C. MHW treatment chambers (n = 5) were heated to about 26°C until sampling 2—‘Induced MHW’, while control chambers (n = 5) remained at ambient conditions of about 21°C. Heating was terminated in the morning of 10 July 2021 and MHW chambers cooled down to ambient temperatures. During the last sampling 3—‘Ambient MHW’, all chambers were at about 24°C—a naturally occurring strong MHW.
Non‐metric multidimensional scaling (nMDS) of the benthic communities based on abundance (panel a) and blotted wet weight biomass (panel b). The Control and marine heatwaves (MHW) treatments are labelled blue and orange, respectively. PERMANOVA results indicate no significate differences between Control and MHW treatments for both abundance (p = 0.44) and biomass (p = 0.12).
Density plots of the size distributions of the most prevalent bioturbators (a) Hediste diversicolor, (b) Macoma balthica and (c) Marenzelleria spp. for treatment (orange) and control (blue). Macoma balthica and Marenzelleria spp. show higher frequencies in smaller sizes in marine heatwaves (MHW) treatment than in control. However, the PERMANOVA test only indicated a significant difference for Marenzelleria spp. (R² = 0.14, pseudo‐F = 12.81, p = 0.001).
Solute fluxes[mmol/m² day⁻¹] (in order from upper to lower panel: Oxygen, ammonium, phosphate, silicate) during night‐time (left column, grey background) and daytime (right column, yellow background) incubations of each sampling day. On day 1. ‘Acclimation’, when all chambers (n = 10) were at same conditions at about 18°C. After 6 days of heating the marine heatwaves (MHW) chambers (n = 5) to about 26°C ‘Induced MHW’ (2.), while control chambers (n = 5) were at about 21°C. After another 6 days, when all chambers were back to natural conditions at about 23°C during an ‘Ambient MHW’ (3.) together with a strong phytoplankton bloom.

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Marine heatwaves amplify benthic community metabolism and solute flux in a seafloor heating experiment
  • Article
  • Full-text available

March 2025

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50 Reads

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1 Citation

Norman Göbeler

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Alf Norkko

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The magnitude and frequency of marine heatwaves are increasing and predicted to intensify, but our ability to understand the real‐world effects on vital benthic ecosystems is lagging behind. Prior insights into the impacts of marine heatwaves are often derived from observational or laboratory studies. Observational studies may not fully disentangle the complexities of potential compound events and typically focus on severe, often lethal marine heatwaves. Laboratory studies, on the contrary, while valuable for understanding specific mechanisms, often use artificial setups and can introduce unnatural disturbances that do not reflect real‐world scenarios. To investigate sublethal temperature effects of marine heatwaves in a natural benthic habitat, we developed a novel approach for inducing elevated water temperatures in situ over several days. The system utilizes domestic underfloor heating technology combined with custom‐made benthic chambers. We placed 10 chambers for 15 days in a bare‐sediment habitat at 2.5 m depth and heated five chambers to 5°C above ambient water temperatures in summer for 6 days, followed by a period of 7 days at ambient temperatures. Incubations during day and night were performed during the experiment to assess changes in ecosystem functioning (solute fluxes) and sediment cores were collected at the end of the experiment to assess the effects of a realistic marine heatwave on benthic community structure. The results indicate that while the benthic community structure remained similar between the treatments, except for a size shift of Marenzelleria spp. towards smaller individuals in the heated treatment, elevated temperatures caused a significant increase in community respiration and amplified the magnitude of either efflux or influx of nutrients (NH4⁺‐N, PO4³⁻‐P and Si). Primary production during daytime incubations remained mostly unaffected by the heatwave treatment, contributing to the concept of heterotrophy being more influenced by increased temperatures than autotrophy. This study confirms the suitability of the novel system for examining the impact of temperature on benthic habitats in situ and demonstrates its potential for the investigation of complex habitats and communities, which are essential for our understanding of the ecosystem‐level effects of climate change. Read the free Plain Language Summary for this article on the Journal blog.

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HOTFLOOR : A benthic chamber system to simulate warming on the seafloor

November 2023

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50 Reads

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1 Citation

Norman Göbeler

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Robin Gottberg

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The frequency of abnormally warm water events is increasing not only in surface waters, but also in subsurface layers, with major impacts on benthic ecosystems. Previous insights on heatwave effects have been obtained through field observations or manipulative laboratory experiments. Here, we introduce a system capable of inducing elevated water temperatures in benthic habitats in situ over several days. The system consists of a commercially available electric boiler, usually applied in domestic underfloor heating, and custom‐designed benthic acrylic glass chambers connected to individual thermostats. Furthermore, the chambers are semi‐open, allowing constant water exchange, maintaining otherwise near‐natural conditions, including oxygen concentrations, while the temperature is elevated. The water exchange can be stopped to facilitate incubations measuring changes in benthic fluxes. We conducted a 15‐d trial study in July 2021 on a bare‐sediment habitat at 2.5 m depth, exposing five chambers to water temperatures 5°C above ambient temperatures for 6 d and comparing with five control chambers. In this assessment, we demonstrate that the temperature control and stability were reliable while maintaining natural oxygen conditions. The modular character of the system permits adaptations for various benthic habitats, facilitating the investigation of elevated temperatures in situ for future climate change scenarios.


Changes in macrofauna bioturbation during repeated heatwaves mediate changes in biogeochemical cycling of nutrients

January 2023

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362 Reads

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10 Citations

The increasing frequency and intensity of marine heatwaves (MHWs) observed worldwide entails changes in the structure and functioning of ecological communities. While severe and extreme heatwaves often have more destructive effects, the more subtle effects of moderate and strong heatwaves may nevertheless affect ecosystem functioning through complex, context-dependent linkages between different processes. Here we conducted a laboratory experiment to study the effects of repeated short-term, strong MHWs on macrofauna bioturbation and associated solute fluxes as a measure of ecosystem functioning using natural soft-sediment communities from the Baltic Sea. Our results showed changes in both bioturbation and biogeochemical cycling of nutrients following short-term, strong heatwaves, which seemed to contribute to an enhanced degradation of organic matter in the seafloor and an enhanced exchange of solutes across the sediment-water interface as well as increased sediment oxygen consumption. Following changes in these processes, the relative contribution of macrofauna and the environmental context to ecosystem functioning was altered. Our results highlight the potential of even shorter-term, strong MHWs of having system-wide impacts due to changes in the mechanistic process of bioturbation underpinning the biogeochemical cycling of nutrients. This study also highlights the need to measure a wide range of variables for a comprehensive understanding of the changes in functioning under disturbances, such as MHWs.


Ninety years of coastal monitoring reveals baseline and extreme ocean temperatures are increasing off the Finnish coast

September 2022

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345 Reads

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23 Citations

Global marine heatwave assessments often rely on satellite-derived sea surface temperature. However, these data have low accuracy in coastal areas, are unable to measure sub-surface temperatures and have only been available since the 1980s. Here, we analyse 90 years of in situ surface and bottom (30 m) water temperature data from a Finnish coastal monitoring site. Water temperatures were significantly higher between 1991–2020 than 1931–1960 and 1961–1990. We find strong differences between satellite-derived and in situ temperatures, with in situ temperatures being lower in autumn and winter and higher in spring. Measurements at the seafloor indicate marine heatwaves occurred during all seasons between 2016 and 2020, with intensities and durations exceeding previous records. Since the 1990s, we find an upward shift of the baseline temperature and increasingly frequent occurrence of temperatures previously considered as an extreme. Our findings highlight the importance of long-term in situ data and choice of climatological reference periods for assessing change.


Literature synthesis of freshwater zooplankton LC50 to Cl⁻ as NaCl. Horizontal lines indicate 95% confidence intervals of reported Cl⁻ LC50 estimates.
Conceptual figure with simulated data showing how population (a), species (b), and subassemblage (c) LC50s were calculated. In all panels, colors denote sites while symbols represent species. In (a), crosses on the x‐axis indicate LC50s, calculated as the Cl⁻ concentration where the population density is predicted to be 50% of the value with no Cl⁻. In (b), species LC50s were calculated as the mean of population estimates, weighted based on model R². The predicted subassemblage LC50 in (c) was calculated for individual sites using a subset of the 19 species found at that site. Note that numbers are provided for illustrative purposes only and do not correspond to observed densities or LC50s. C.I., confidence interval; ind., individual; pop., populations; sp., species.
Cl⁻ tolerance (LC50) of freshwater zooplankton species occurring at more than one site in the coordinated mesocosm experiment. Species are grouped by major taxon (top = copepods; middle = cladocerans; bottom = rotifers). Colored circles and lines represent site‐specific population LC50 ± 95% confidence intervals. Colors correspond to sites in Fig. 5. The size of the circles represents the confidence in the model predicting the population LC50 (GAM R²). The number of sites at which each species is found is shown in parentheses. The bolded black line and black circle are the weighted mean ± standard error (95% confidence intervals) representing species‐LC50s when averaging population‐LC50s across sites.
Predictors of intraspecific variation in Cl⁻ tolerance. Each circle represents a species listed in Fig. 3. A regression line indicates the relationship between intraspecific variation and the one significant predictor of intraspecific variation (community dissimilarity). Cla., cladocerans; com., community; Cop., copepods; env., environmental; Rot., rotifers; s, standard deviation.
Community‐weighted mean Cl⁻ tolerance (subassemblage‐LC50) of freshwater zooplankton from each site (squares) where at least 3 of the 19 species were detected. Simulated‐LC50s were drawn from species' LC50 mean and standard deviation excluding the focal population to test prediction accuracy when intraspecific variation was not considered. Simulated LC50s are shown using colored dots and boxplots. Asterisks indicate subassemblage‐LC50s that deviate significantly from simulated‐LC50s at p < 0.05 (note that p‐values were adjusted for multiple comparisons using the false discovery rate method). The numbers in parentheses represent the number of species used to calculate the subassemblage‐LC50s.
Widespread variation in salt tolerance within freshwater zooplankton species reduces the predictability of community‐level salt tolerance

August 2022

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411 Reads

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16 Citations

The salinization of freshwaters is a global threat to aquatic biodiversity. We quantified variation in chloride (Cl−) tolerance of 19 freshwater zooplankton species in four countries to answer three questions: (1) How much variation in Cl− tolerance is present among populations? (2) What factors predict intraspecific variation in Cl− tolerance? (3) Must we account for intraspecific variation to accurately predict community Cl− tolerance? We conducted field mesocosm experiments at 16 sites and compiled acute LC50s from published laboratory studies. We found high variation in LC50s for Cl− tolerance in multiple species, which, in the experiment, was only explained by zooplankton community composition. Variation in species‐LC50 was high enough that at 45% of lakes, community response was not predictable based on species tolerances measured at other sites. This suggests that water quality guidelines should be based on multiple populations and communities to account for large intraspecific variation in Cl− tolerance.


Fig. 1. GAMs and 95% CIs (shaded regions) for cladoceran zooplankton abundance (black line) and chlorophyll a concentration (i.e., phytoplankton biomass; green line) among 16 experimental sites throughout North America and Europe. Cladocerans are featured as they are generally regarded as the largest, highest-biomass filter-feeding zooplankton grazers in our study systems. The vertical red dashed line indicates the lethal concentration value (LC 50 ) representing the Cl À concentration at which there was a 50% reduction in abundance of each zooplankton taxa for each site with the associated 95% CIs (red shaded region around the dashed line). Graphs for calanoid and cyclopoid copepods and rotifers are shown in SI Appendix, Appendix 4, Figs. 4.2-4.4. No cladocerans were found at the Tavernoles study site in Spain.
Fig. 2. LC 50 representing the Cl À concentration at which there was a 50% reduction in abundance of each zooplankton taxa across experimental sites (A) Cladocerans, (B) Calanoids, (C) Cyclopoids, and (D) Rotifers. The horizontal red lines highlight the lowest global Cl À thresholds for protecting aquatic life, which are set by governments in Canada (solid line, 120 mg Cl À /L) and the United States (dashed line, 230 mg Cl À /L). Vertical bars represent 95% CIs. Points and 95% CIs are red when they overlap with or were below the US threshold of 230 mg Cl À /L. When a zooplankton taxon is not found in the mesocosm experiment, it is noted as "NA." One experiment did not count rotifers, denoted with an asterisk. LC 50 values were
Fig. 3. Mean (±1 SE) of the estimated percent change in zooplankton abundances for each taxa at the Canadian (120 mg Cl À /L) and US (230 mg Cl À /L) chronic threshold for Cl À . Percent changes were estimated using GAM models to compare the predicted abundances at each threshold to the predicted abundance in control conditions.
Current water quality guidelines across North America and Europe do not protect lakes from salinization

March 2022

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544 Reads

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93 Citations

Proceedings of the National Academy of Sciences

Significance The salinity of freshwater ecosystems is increasing worldwide. Given that most freshwater organisms have no recent evolutionary history with high salinity, we expect them to have a low tolerance to elevated salinity caused by road deicing salts, agricultural practices, mining operations, and climate change. Leveraging the results from a network of experiments conducted across North America and Europe, we showed that salt pollution triggers a massive loss of important zooplankton taxa, which led to increased phytoplankton biomass at many study sites. We conclude that current water quality guidelines established by governments in North America and Europe do not adequately protect lake food webs, indicating an immediate need to establish guidelines where they do not exist and to reassess existing guidelines.


Variation in (a–c) crustacean and (d–f) rotifer community abundances and diversity metrics in response to chloride additions across lake experiments. Each colored line distinctly represents one of the source lakes (individual lake linear regression models), while thicker black lines represent the general trend across lakes (LMM; significant models only, slope coefficients provided). For among‐lake LMMs, both marginal and conditional R² are provided (Rm² and Rc², respectively). See Table S3 for parameter estimates.
Taxon‐level abundance responses of (a) crustaceans and (b) rotifers to chloride additions across lake experiments. Color saturation represents lake‐specific slopes of models, reflecting taxon‐level sensitivity to chloride. Unframed, gray squares represent nonsignificant relationships whereas white areas indicate the absence of taxa in each lake. Abundance data are log10‐transformed. See Table S5 for parameter estimates.
Variation in crustacean (a) trait assemblages and (b, c) functional diversity in response to chloride additions across lake experiments. (a) For each lake, fourth corner coefficients show the strength of association between traits and chloride concentrations, from positive (blue) to negative (red). See Table S5 for parameter estimates. (b, c) Each colored line distinctly represents one of the source lakes (individual lake linear regression model; colors as in Fig. 1), while thicker black lines represent the general trend across lakes (LMMs, slope coefficients provided). For among‐lake LMMs, both marginal and conditional R² are provided (Rm² and Rc², respectively). See Tables S3 and S4 for parameter estimates.
Comparisons of chloride sensitivity among (a–c) crustacean and (d–f) rotifer community‐level measures (abundance and diversity metrics) and across source lakes. (a, d) Chloride concentrations correspond to 50% reductions in community‐level measures (analogous to half maximal effective concentration EC50), and (b, c, e, f) specific proportions (%) of reduction are shown at Canadian (120 mg Cl⁻ L⁻¹, middle column) and U.S. (230 mg Cl⁻ L⁻¹, right column) water quality guidelines. Each colored point distinctly represents one of the source lakes (individual lake linear model; colors as in Figs. 1 and 3); large transparent circles illustrate the overall value across lakes (among‐lake LMMs). Only values from significant models are shown.
Lake salinization drives consistent losses of zooplankton abundance and diversity across coordinated mesocosm experiments

February 2022

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515 Reads

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46 Citations

Human-induced salinization increasingly threatens inland waters; yet we know little about the multifaceted response of lake communities to salt contamination. By conducting a coordinated mesocosm experiment of lake salinization across 16 sites in North America and Europe, we quantified the response of zooplankton abundance and (taxonomic and functional) community structure to a broad gradient of environmentally relevant chloride concentrations, ranging from 4 to ca. 1400 mg Cl/L. We found that crustaceans were distinctly more sensitive to elevated chloride than rotifers; yet, rotifers did not show compensatory abundance increases in response to crustacean declines. For crustaceans, our among-site comparisons indicate: (1) highly consistent decreases in abundance and taxon richness with salinity; (2) widespread chloride sensitivity across major taxonomic groups (Cladocera, Cyclopoida, and Calanoida); and (3) weaker loss of functional than taxonomic diversity. Overall, our study demonstrates that aggregate properties of zooplankton communities can be adversely affected at chloride concentrations relevant to anthropogenic salinization in lakes.


Anti-Fouling Effects of Saponin-Containing Crude Extracts from Tropical Indo-Pacific Sea Cucumbers

March 2020

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271 Reads

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16 Citations

Sea cucumbers are bottom dwelling invertebrates, which are mostly found on subtropical and tropical sea grass beds, sandy reef flats, or reef slopes. Although constantly exposed to fouling communities in these habitats, many species are surprisingly free of invertebrate epibionts and microfouling algae such as diatoms. In our study, we investigated the anti-fouling (AF) activities of different crude extracts of tropical Indo-Pacific sea cucumber species against the fouling diatom Cylindrotheca closterium. Nine sea cucumber species from three genera (i.e., Holothuria, Bohadschia, Actinopyga) were selected and extracted to assess their AF activities. To verify whether the sea cucumber characteristic triterpene glycosides were responsible for the observed potent AF activities, we tested purified fractions enriched in saponins isolated from Bohadschia argus, representing one of the most active anti-fouling extracts. Saponins were quantified by vanillin-sulfuric acid colorimetric assays and identified by LC-MS and LC-MS/MS analyses. We were able to demonstrate that AF activities in sea cucumber extracts were species-specific, and growth inhibition as well as attachment of the diatom to surfaces is dependent on the saponin concentration (i.e., Actinopyga contained the highest quantities), as well as on the molecular composition and structure of the present saponins (i.e., Bivittoside D derivative was the most bioactive compound). In conclusion, the here performed AF assay represents a promising and fast method for selecting the most promising bioactive organism as well as for identifying novel compounds with potent AF activities for the discovery of potentially novel pharmacologically active natural products.

Citations (8)


... Cascading effects in food webs and ecosystem regime shifts have been studied and observed in the open Baltic Sea, but less so in coastal areas (Yletyinen et al. 2016;Tomczak et al. 2022). Our understanding of degrading ecosystem functioning is still lagging, but experimental in situ testing of stressors such as hypoxia (e.g., Villnäs et al. 2013) or marine heatwaves (Göbeler et al. 2025) suggest profound changes to, e.g., nutrient cycling. More attention should also be paid to less studied parts of ecosystems, e.g., microbial food webs and sea ice communities (Viitasalo and Bonsdorff 2022). ...

Reference:

Marine biodiversity loss in Finnish coastal waters: Evidence and implications for management
Marine heatwaves amplify benthic community metabolism and solute flux in a seafloor heating experiment

... Ecosystem functioning of benthic communities in terms of nutrient cycling is highly complex and determined by various biotic and abiotic factors . In this study, relatively natural oxygen conditions inside the chambers were maintained through the water exchange with the surrounding water for the majority of the time (unless incubating) and followed the ambient conditions (see Göbeler et al., 2023 for details). The three-way PERMANOVA revealed a significant effect of the Sampling occasion and Treatment on ecosystem functioning, while Sampling accounted for the majority of the variance. ...

HOTFLOOR : A benthic chamber system to simulate warming on the seafloor
  • Citing Article
  • November 2023

... While the general importance of temperature for biochemical processes is well known, only a few studies have thus far investigated the effects of episodic events, such as MHWs on nutrient fluxes. In a laboratory study with intact sediment cores from a deeper, muddy site, Kauppi et al. (2023) found that repeated, strong MHWs can increase bioturbation activities and thus enhance carbon remineralization. While a similar effect was detected by Kauppi and Villnäs (2022) also utilizing intact sediment cores from a muddy site, this study also investigated even higher temperatures that resulted in a severe reduction in oxygen consumption possibly related to the inactivation of microbial processes. ...

Changes in macrofauna bioturbation during repeated heatwaves mediate changes in biogeochemical cycling of nutrients

... Understanding these complex interactions is crucial for predicting and mitigating the long-term ecological consequences of multiple pressures (Reckermann et al. 2022;Viitasalo and Bonsdorff 2022). Moreover, the impact of the increasing prevalence of marine heatwaves are virtually unknown (Goebeler et al. 2022) and considering the high use of the coastal areas in recreation (i.e., summer cottages, boating) more studies are also needed on the cumulative biodiversity effects of local habitat disturbances . ...

Ninety years of coastal monitoring reveals baseline and extreme ocean temperatures are increasing off the Finnish coast

... Another factor that may strongly influence NaCl toxicity for organisms embedded in their natural ecosystem is a previous history of salt exposure. Researchers have uncovered evidence of intraspecific variation in NaCl tolerance in a number of taxonomic groups including yeast (Bell & Gonzalez, 2011), amphibians (Albecker et al., 2021;Brady, 2012Brady, , 2013, fish (Spence et al., 2012), and zooplankton (Arnott et al., 2023;Latta et al., 2012;Liu & Steiner, 2017;Loureiro et al., 2012;Venâncio et al., 2018). In fact, a recent study demonstrated that intraspecific variation in the chloride tolerance of zooplankton species was so high that community level responses to NaCl exposure could not be predicted based on established species level toxicity thresholds (Arnott et al., 2023). ...

Widespread variation in salt tolerance within freshwater zooplankton species reduces the predictability of community‐level salt tolerance

... and can disturb nutrient and energy flow at the ecosystem level (Cochero et al., 2017;Hintz & Relyea, 2019;Hintz, Arnott et al., 2022). ...

Current water quality guidelines across North America and Europe do not protect lakes from salinization

Proceedings of the National Academy of Sciences

... The SITES AquaNet mesocosm infrastructure is another network of mesocosms located in five Swedish lakes, covering a 760 km latitudinal gradient (Urrutia-Cordero et al. 2021b). A cross-continental lake salinisation experiment incorporated three of the SITES AquaNet facilities, together with fifteen other existing mesocosm systems across Europe and North America, to study global patterns of salinisation on zooplankton communities (Hébert et al. 2022). While it was less standardised in terms of the system (many different mesocosm systems were used, ranging from 20-32 experimental units and 80-2500 l volumes), it was still highly standardised in terms of salinity treatment (an unreplicated regression design), study time frame, and sampling protocol. ...

Lake salinization drives consistent losses of zooplankton abundance and diversity across coordinated mesocosm experiments

... [13][14][15] Notwithstanding, ASP VI belongs to Biopharmaceutics Classification System class III (high solubility/low permeability) with poor oral bioavailability (<0.13%) and strong hydrophilic properties. These properties are attributed to its physical properties such as high molecular weight (>500 Da) and a large number of hydrogen bonds (>12), 16 exhibiting amphipathic properties that easily form the self-assembled structures. Numerous studies have shown that the active ingredients with different amounts of hydroxyl and carboxyl groups could spontaneously self-assemble into nanomicelles through non-covalent interactions such as π-π stacking, electrostatic interactions, hydrogen bonding and coordination to change its mucosal permeability. ...

Anti-Fouling Effects of Saponin-Containing Crude Extracts from Tropical Indo-Pacific Sea Cucumbers