Long-term declines in chlorophyll a and variable phenology revealed by a 60-year estuarine plankton time series.

Long-term ecological time series provide a unique perspective on the emergent properties of ecosystems. In aquatic systems, phytoplankton form the base of the food web and their biomass, measured as the concentration of the photosynthetic pigment chlorophyll a (chl a), is an indicator of ecosystem quality. We analyzed temporal trends in chl a from the Long-Term Plankton Time Series in Narragansett Bay, Rhode Island, USA, a temperate estuary experiencing long-term warming and changing anthropogenic nutrient inputs. Dynamic linear models were used to impute and model environmental variables (1959 to 2019) and chl a concentrations (1968 to 2019). A long-term chl a decrease was observed with an average decline in the cumulative annual chl a concentration of 49% and a marked decline of 57% in winter-spring bloom magnitude. The long-term decline in chl a concentration was directly and indirectly associated with multiple environmental factors that are impacted by climate change (e.g., warming temperatures, water column stratification, reduced nutrient concentrations) indicating the importance of accounting for regional climate change effects in ecosystem-based management. Analysis of seasonal phenology revealed that the winter-spring bloom occurred earlier, at a rate of 4.9 ± 2.8 d decade-1. Finally, the high degree of temporal variation in phytoplankton biomass observed in Narragansett Bay appears common among estuaries, coasts, and open oceans. The commonality among these marine ecosystems highlights the need to maintain a robust set of phytoplankton time series in the coming decades to improve signal-to-noise ratios and identify trends in these highly variable environments.

Synergistic digestibility effect by planktonic natural food and habitat renders high digestion efficiency in agastric aquatic consumers.

A digestibility enhancing effect of natural food on stomachless fish model (Cyprinus carpio) was verified by fluorogenic substrate assays of enzymatic activities in experimental pond carp gut flush and planktonic food over a full vegetative season. Then compared with size-matched conspecific grown artificially (tank carp) and an advanced omnivore species possessing true stomach (tilapia, Oreochromis niloticus). Results suggested activities of digestive enzymes (except amylolytic) were significantly higher in pond carp (p ≤ 0.05) than in the size-matched tank carp. Even compared to tilapia, pond carp appeared superior (p < 0.05; proteolytic or chitinolytic activities) or comparable (p > 0.05; phosphatase or cellulolytic activities). Amylolytic, chitinolytic, and phosphatases activities in pond carp gut significantly increased (p ≤ 0.01) over season. Several orders-of-magnitude higher enzymatic activities were detected in planktonic natural food than expressed in carp gut. Amino acid markers in planktonic food revealed a higher share of zooplankton (microcrustaceans), but not phytoplankton, synchronized with higher activities of complex polysaccharide-splitting enzymes (cellulolytic and chitinolytic) in fish gut. Periods of clear water phase low in chlorophyll-a and nutrients, but high in certain zooplankton (preferably cladocerans), may create a synergistic digestibility effect in pond carp. We conclude aquatic ecosystem components (natural food, water, microbiota) enhance fishes' hydrolyzing capabilities of C/N/P macromolecules and even their complex polymers such as cellulose, chitin, and maybe phytate (to be validated), to the extent that being stomachless is not an issue. Aquatic nutritional ecologists may consider that laboratory-based understandings of digestibility may underestimate digestion efficiency of free-ranging fish in ponds or lakes.

A Lagrangian model for drifting ecosystems reveals heterogeneity-driven enhancement of marine plankton blooms.

Marine plankton play a crucial role in carbon storage, global climate, and ecosystem function. Planktonic ecosystems are embedded in patches of water that are continuously moving, stretching, and diluting. These processes drive inhomegeneities on a range of scales, with implications for the integrated ecosystem properties, but are hard to characterize. We present a theoretical framework that accounts for all these aspects; tracking the water patch hosting a drifting ecosystem along with its physical, environmental, and biochemical features. The theory resolves patch dilution and internal physical mixing as a function of oceanic strain and diffusion. Ecological dynamics are parameterized by an idealized nutrient and phytoplankton population and we specifically capture the time evolution of the biochemical spatial variances to represent within-patch heterogeneity. We find that, depending only on the physical processes to which the water patch is subjected, the plankton biomass response to a resource perturbation can vary in size up to six times. This work indicates that we must account for these processes when interpreting and modeling marine ecosystems and provides a framework with which to do so.

Multiyear analysis uncovers coordinated seasonality in stocks and composition of the planktonic food web in the Baltic Sea proper.

The planktonic realm from bacteria to zooplankton provides the baseline for pelagic aquatic food webs. However, multiple trophic levels are seldomly included in time series studies, hampering a holistic understanding of the influence of seasonal dynamics and species interactions on food web structure and biogeochemical cycles. Here, we investigated plankton community composition, focusing on bacterio-, phyto- and large mesozooplankton, and how biotic and abiotic factors correlate at the Linnaeus Microbial Observatory (LMO) station in the Baltic Sea from 2011 to 2018. Plankton communities structures showed pronounced dynamic shifts with recurring patterns. Summarizing the parts of the planktonic microbial food web studied here to total carbon, a picture emerges with phytoplankton consistently contributing > 39% while bacterio- and large mesozooplankton contributed ~ 30% and ~ 7%, respectively, during summer. Cyanophyceae, Actinobacteria, Bacteroidetes, and Proteobacteria were important groups among the prokaryotes. Importantly, Dinophyceae, and not Bacillariophyceae, dominated the autotrophic spring bloom whereas Litostomatea (ciliates) and Appendicularia contributed significantly to the consumer entities together with the more traditionally observed mesozooplankton, Copepoda and Cladocera. Our findings of seasonality in both plankton composition and carbon stocks emphasize the importance of time series analyses of food web structure for characterizing the regulation of biogeochemical cycles and appropriately constraining ecosystem models.

Staphylococci planktonic and biofilm environments differentially affect osteoclast formation.

INTRODUCTION: The pathophysiology of chronic implant-related bone infections is characterized by an increase in osteoclast numbers and enhanced bone resorption. Biofilms are a major reason for chronicity of such infections as the biofilm matrix protects bacteria against antibiotics and impairs the function of immune cells. Macrophages are osteoclast precursor cells and therefore linked to inflammation and bone destruction. OBJECTIVE AND METHOD: Investigations on the impact of biofilms on the ability of macrophages to form osteoclasts are yet missing and we, therefore, analyzed the effect of Staphylococcus aureus (SA) and Staphylococcus epidermidis (SE) planktonic and biofilm environments on osteoclastogenesis using RAW 264.7 cells and conditioned media (CM). RESULTS: Priming with the osteoclastogenic cytokine RANKL before CM addition enabled the cells to differentiate into osteoclasts. This effect was highest in SE planktonic or SA biofilm CM. Simultaneous stimulation with CM and RANKL, however, suppressed osteoclast formation and resulted in formation of inflammation-associated multinucleated giant cells (MGCs) which was most pronounced in SE planktonic CM. CONCLUSION: Our data indicate that the biofilm environment and its high lactate levels are not actively promoting osteoclastogenesis. Hence, the inflammatory immune response against planktonic bacterial factors through Toll-like receptors seems to be the central cause for the pathological osteoclast formation. Therefore, immune stimulation or approaches that aim at biofilm disruption need to consider that this might result in enhanced inflammation-mediated bone destruction.

Direct input of monitoring data into a mechanistic ecological model as a way to identify the phytoplankton growth-rate response to temperature variations.

We present an approach (knowledge-and-data-driven, KDD, modeling) that allows us to get closer to understanding the processes that affect the dynamics of plankton communities. This approach, based on the use of time series obtained as a result of ecosystem monitoring, combines the key features of both the knowledge-driven modeling (mechanistic models) and data-driven (DD) modeling. Using a KDD model, we reveal the phytoplankton growth-rate fluctuations in the ecosystem of the Naroch Lakes and determine the degree of phase synchronization between fluctuations in the phytoplankton growth rate and temperature variations. More specifically, we estimate a numerical value of the phase locking index (PLI), which allows us to assess how temperature fluctuations affect the dynamics of phytoplankton growth rates. Since, within the framework of KDD modeling, we directly include the time series obtained as a result of field measurements in the model equations, the dynamics of the phytoplankton growth rate obtained from the KDD model reflect the behavior of the lake ecosystem as a whole, and PLI can be considered as a holistic parameter.

Impact of climate warming on phenological asynchrony of plankton dynamics across Europe.

Climate warming alters the seasonal timing of biological events. This raises concerns that species-specific responses to warming may de-synchronize co-evolved consumer-resource phenologies, resulting in trophic mismatch and altered ecosystem dynamics. We explored the effects of warming on the synchrony of two events: the onset of the phytoplankton spring bloom and the spring/summer maximum of the grazer Daphnia. Simulation of 16 lake types over 31 years at 1907 North African and European locations under 5 climate scenarios revealed that the current median phenological delay between the two events varies greatly (20-190 days) across lake types and geographic locations. Warming moves both events forward in time and can lengthen or shorten the delay between them by up to ±60 days. Our simulations suggest large geographic and lake-specific variations in phenological synchrony, provide quantitative predictions of its dependence on physical lake properties and geographic location and highlight research needs concerning its ecological consequences.

The Mixoplankton Database (MDB): Diversity of photo-phago-trophic plankton in form, function, and distribution across the global ocean.

Protist plankton are major members of open-water marine food webs. Traditionally divided between phototrophic phytoplankton and phagotrophic zooplankton, recent research shows many actually combine phototrophy and phagotrophy in the one cell; these protists are the "mixoplankton." Under the mixoplankton paradigm, "phytoplankton" are incapable of phagotrophy (diatoms being exemplars), while "zooplankton" are incapable of phototrophy. This revision restructures marine food webs, from regional to global levels. Here, we present the first comprehensive database of marine mixoplankton, bringing together extant knowledge of the identity, allometry, physiology, and trophic interactivity of these organisms. This mixoplankton database (MDB) will aid researchers that confront difficulties in characterizing life traits of protist plankton, and it will benefit modelers needing to better appreciate ecology of these organisms with their complex functional and allometric predator-prey interactions. The MDB also identifies knowledge gaps, including the need to better understand, for different mixoplankton functional types, sources of nutrition (use of nitrate, prey types, and nutritional states), and to obtain vital rates (e.g. growth, photosynthesis, ingestion, factors affecting photo' vs. phago' -trophy). It is now possible to revisit and re-classify protistan "phytoplankton" and "zooplankton" in extant databases of plankton life forms so as to clarify their roles in marine ecosystems.

Increasing diversity and biotic homogenization of lake plankton during recovery from acidification.

Determining biodiversity responses to environmental change, such as acidification, is critical for ecosystem projections under future global change scenarios. Here, we analyzed three plankton communities of phytoplankton, crustaceans and rotifers in 28 lakes in the Adirondack Park, USA, during 1994-2012, and examined the spatiotemporal trends in their alpha and beta diversity during recovery from acidification. For all plankton assemblages, Shannon diversity increased towards recent years and high lake pH, and there was an increasing community dissimilarity with pH changes. The spatial mean Bray-Curtis dissimilarities across all lakes decreased over time for phytoplankton and rotifers leading to an increase in spatial homogenization. Such a homogenization cooccurred however with the overall increasing diversity in this region, which contrasts with the previous classic view that homogenization is mainly driven by loss of species and results in biodiversity loss. We further observed lower temporal mean beta diversity in low-pH lakes for crustaceans and rotifers, but not for phytoplankton. Generally, spatial and temporal mean beta diversity of the three taxonomic groups were primarily driven by lake-water ion variables, and rotifers were also constrained by nutrients and climate. Collectively, our results show how and why plankton community compositions vary over space along with acidification recovery, and further highlight the importance of spatiotemporal studies combined with long-term monitoring programs in assessing biodiversity change during the recovery of disturbed ecosystems.

Planktonic ciliate community driven by environmental variables and cyanobacterial blooms: A 9-year study in two subtropical reservoirs.

It is well-established that environmental variability and cyanobacterial blooms have major effects on the assembly and functioning of bacterial communities in both marine and freshwater habitats. It remains unclear, however, how the ciliate community responds to such changes over the long-term, particularly in subtropical lake and reservoir ecosystems. We analysed 9-year planktonic ciliate data series from the surface water of two subtropical reservoirs to elucidate the role of cyanobacterial bloom and environmental variabilities on the ciliate temporal dynamics. We identified five distinct periods of cyanobacterial succession in both reservoirs. Using multiple time-scale analyses, we found that the interannual variability of ciliate communities was more strongly related to cyanobacterial blooms than to other environmental variables or to seasonality. Moreover, the percentage of species turnover across cyanobacterial bloom and non-bloom periods increased significantly with time over the 9-year period. Phylogenetic analyses further indicated that 84 %-86 % of ciliate community turnover was governed by stochastic dispersal limitation or undominated processes, suggesting that the ciliate communities in subtropical reservoirs were mainly controlled by neutral processes. However, short-term blooms increased the selection pressure and drove 30 %-53 % of the ciliate community turnover. We found that the ciliate community composition was influenced by environmental conditions with nutrients, cyanobacterial biomass and microzooplankton having direct and/or indirect significant effects on the ciliate taxonomic or functional community dynamics. Our results provide new insights into the long-term temporal dynamics of planktonic ciliate communities under cyanobacterial bloom disturbance.

Factors regulating phytoplankton biomass along the Indian coast: elucidation with long-term data.

Long-term variations in phytoplankton biomass from nine coastal states along the Indian coast were studied and co-related with biotic and abiotic factors. Surface water temperature, dissolved inorganic nutrients (nitrogen and phosphorous), and plankton (phytoplankton and zooplankton) biomass data were collected between 1992 and 2015. Linear regression analysis showed a considerable increase in dissolved inorganic nutrients. A substantial increase in the frequency and intensity of phytoplankton blooms (> 3 mg/m3) was observed along the Indian coast. The considerable increase in dissolved inorganic nutrient concentrations could be the major reason for an increase in phytoplankton bloom occurrences. Availability of light and periodicity in zooplankton population were also regulating phytoplankton biomass. Our results revealed that multiple factors are influencing phytoplankton biomass along the Indian coast; particularly, the increase in nutrient concentrations is promoting plankton biomass. Moreover, significant increase in zooplankton biomass can have substantial impact on the biogeochemical cycling and energy transfer to higher trophic levels.

Farnesol modulation of Rhodotorula mucilaginosa in biofilm and planktonic forms.

Biofilms are important to the virulence of human pathogenic fungi, and some molecules have been found to play key roles in the growth and regulation of fungal biofilms. Farnesol, one of these molecules, is well-described for some microorganisms but is still scarcely known for Rhodotorula spp. This study aimed to evaluate the influence of farnesol on the biofilm of R. mucilaginosa. Initially, screening with 0.2 mM to 2.1 mM of farnesol was evaluated against planktonic forms. A concentration of this compound was then chosen and evaluated for its effect on biofilm in formation and on preformed biofilm after 24, 48 and 72 hours. The impact of farnesol was evaluated by colony-forming units (CFU) counts, determination of metabolic activity and quantification of total biomass. In the presence of 0.9 mM, farnesol was able to decrease the CFU number, at 48 hours, when the biofilm was in formation, although it did not affect the preformed biofilms. Thus, our results show that farnesol exerts a modulating activity during biofilm formation for R. mucilaginosa, with this compound reducing the metabolic activity and total biomass of the biofilms.

Beyond the fish-Daphnia paradigm: testing the potential for pygmy backswimmers (Neoplea striola) to cause trophic cascades in subtropical ponds.

Trophic cascades, or indirect effects of predators on non-adjacent lower trophic levels, are a classic phenomenon in ecology, and are thought to be strongest in aquatic ecosystems. Most research on freshwater trophic cascades focused on temperate lakes, where fish are present and where Daphnia frequently dominate the zooplankton community. These studies identified that Daphnia often play a key role in facilitating trophic cascades by linking fish to algae with strong food web interactions. However, Daphnia are rare or absent in most tropical and subtropical lowland freshwaters, and fish are absent from small and temporary water bodies, where invertebrates fill the role of top predator. While invertebrate predators are ubiquitous in freshwater systems, most have received little attention in food web research. Therefore, we aimed to test whether trophic cascades are possible in small warmwater ponds where Daphnia are absent and small invertebrates are the top predators. We collected naturally occurring plankton communities from small fishless water bodies in central Texas and propagated them in replicate pond mesocosms. We removed zooplankton from some mesocosms, left the plankton community intact in others, and added one of two densities of the predaceous insect Neoplea striola to others. Following an incubation period, we then compared biomasses of plankton groups to assess food web effects between the trophic levels, including whether Neoplea caused a trophic cascade by reducing zooplankton. The zooplankton community became dominated by copepods which prefer large phytoplankton and exhibit a fast escape response. Perhaps due to these qualities of the copepods and perhaps due to other reasons such as high turbidity impairing predation, no evidence for food web effects were found other than somewhat weak evidence for zooplankton reducing large phytoplankton. More research is needed to understand the behavior and ecology of Neoplea, but trophic cascades may generally be weak or absent in fishless low latitude lowland water bodies where Daphnia are rare.

Evaluating the effects of aquaculture on the freshwater lake from the perspective of plankton communities: The diversity, co-occurrence patterns and their underlying mechanisms.

Aquaculture has significant impacts on freshwater lakes, but plankton communities, as key components of the microbial food web, are rarely considered when assessing the impacts of aquaculture. Revealing the dynamics of plankton communities, including bacterioplankton, phytoplankton and zooplankton, under anthropological disturbances is critical for predicting the freshwater ecosystem functioning in response to future environmental changes. In the present study, we examined the impacts of aquaculture on water quality, plankton diversity and the co-occurrence patterns within plankton metacommunities in a shallow freshwater lake. The study zones are influenced by the 20-year historical intensive aquaculture, but now they are undergoing either ecological aquaculture or ecological restoration. Our results showed that ecological aquaculture was more efficient in nitrogen removal than ecological restoration. Moreover, lower bacterioplankton diversity but higher phytoplankton and zooplankton diversity were found in the ecological aquaculture and ecological restoration zones compared to the control zone. The lower network connectivity of the plankton metacommunities in the ecological aquaculture and ecological restoration zones indicated the decreasing complexity of potential microbial food web, suggesting a possible lower resistance of the plankton metacommunities to future disturbance. Furthermore, plankton communities of different trophic levels were driven under distinct mechanisms. The bacterioplankton community was primarily affected by abiotic factors, whereas the phytoplankton and zooplankton communities were explained more by trophic interactions. These results revealed the impacts of aquaculture on the plankton communities and their potential interactions, thereby providing fundamental information for better understanding the impacts of aquaculture on freshwater ecosystem functioning.

The dynamic trophic architecture of open-ocean protist communities revealed through machine-guided metatranscriptomics.

Intricate networks of single-celled eukaryotes (protists) dominate carbon flow in the ocean. Their growth, demise, and interactions with other microorganisms drive the fluxes of biogeochemical elements through marine ecosystems. Mixotrophic protists are capable of both photosynthesis and ingestion of prey and are dominant components of open-ocean planktonic communities. Yet the role of mixotrophs in elemental cycling is obscured by their capacity to act as primary producers or heterotrophic consumers depending on factors that remain largely uncharacterized. Here, we develop and apply a machine learning model that predicts the in situ trophic mode of aquatic protists based on their patterns of gene expression. This approach leverages a public collection of protist transcriptomes as a training set to identify a subset of gene families whose transcriptional profiles predict trophic mode. We applied our model to nearly 100 metatranscriptomes obtained during two oceanographic cruises in the North Pacific and found community-level and population-specific evidence that abundant open-ocean mixotrophic populations shift their predominant mode of nutrient and carbon acquisition in response to natural gradients in nutrient supply and sea surface temperature. Metatranscriptomic data from ship-board incubation experiments revealed that abundant mixotrophic prymnesiophytes from the oligotrophic North Pacific subtropical gyre rapidly remodeled their transcriptome to enhance photosynthesis when supplied with limiting nutrients. Coupling this approach with experiments designed to reveal the mechanisms driving mixotroph physiology provides an avenue toward understanding the ecology of mixotrophy in the natural environment.

Abrupt conclusion of the late Miocene-early Pliocene biogenic bloom at 4.6-4.4 Ma.

The late Miocene-early Pliocene biogenic bloom was an extended time interval characterised by elevated ocean export productivity at numerous locations. As primary productivity is nutrient-limited at low-to-mid latitudes, this bloom has been attributed to an increase or a redistribution of available nutrients, potentially involving ocean-gateway or monsoon-related mechanisms. While the exact causal feedbacks remain debated, there is even less consensus on what caused the end of the biogenic bloom. Here, we compile Mio-Pliocene paleoproductivity proxy data from all major ocean basins to evaluate the timing and pacing of this termination. This systematic analysis reveals an abrupt and sustained reduction in low-latitude ocean productivity at 4.6-4.4 Ma. The decline in productivity coincided with a prolonged period of low orbital eccentricity and a shift towards lower-amplitude obliquity, an astronomical configuration linked to reduced East Asian Monsoon intensity and decreased riverine nutrient supply.

Exploring the impact of climate change on the global distribution of non-spinose planktonic foraminifera using a trait-based ecosystem model.

Planktonic foraminifera are one of the primary calcifiers in the modern ocean, contributing 23%-56% of total global pelagic carbonate production. However, a mechanistic understanding of how physiology and environmental conditions control their abundance and distribution is lacking, hindering the projection of the impact of future climate change. This understanding is important, not only for ecosystem dynamics, but also for marine carbon cycling because of foraminifera's key role in carbonate production. Here we present and apply a global trait-based ecosystem model of non-spinose planktonic foraminifera ('ForamEcoGEnIE') to assess their ecology and global distribution under future climate change. ForamEcoGEnIE considers the traits of calcium carbonate production, shell size, and foraging. It captures the main characteristic of biogeographical patterns of non-spinose species - with maximum biomass concentrations found in mid- to high-latitude waters and upwelling areas. The model also reproduces the magnitude of global carbonate production relatively well, although the foraminifera standing stock is systematically overestimated. In response to future scenarios of rising atmospheric CO2 (RCP6 and RCP8.5), on a regional scale, the modelled foraminifera biomass and export flux increases in the subpolar regions of the North Atlantic and the Southern Ocean while it decreases everywhere else. In the absence of adaptation, the biomass decline in the low-latitude South Pacific suggests extirpation. The model projects a global average loss in non-spinose foraminifera biomass between 8% (RCP6) and 11% (RCP8.5) by 2050 and between 14% and 18% by 2100 as a response to ocean warming and associated changes in primary production and ecological dynamics. Global calcium carbonate flux associated with non-spinose foraminifera declines by 13%-18% by 2100. That decline can slow down the ocean carbonate pump and create short-term positive feedback on rising atmospheric pCO2 .