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Satellite observations have given us a clear idea of the changes in chlorophyll in the surface ocean on both a seasonal and interannual basis, but repeated observations at depth are much rarer. The permanently-stratified subtropical gyres in the Atlantic are highly oligotrophic, with most production centred on a deep chlorophyll maximum (DCM) just above the nitracline. This study explores the variations in this feature in the core of both gyres, considering both seasonal and interannual variations, and the linkages between changes at the surface and sub-surface. The in situ observations come from the Atlantic Meridional Transect (AMT), a long-running UK monitoring programme, and also from biogeochemical Argo floats. AMT provides measurements spanning more than 25 years directed through the centres of these gyres, but samples only 2 to 4 months per year and thus cannot resolve the seasonal variations, whereas the profiling floats give coverage throughout the year, but without the rigid spatial repeatability. These observational records are contrasted with representation of the centres of the gyres in two different biogeochemical models: MEDUSA and ERSEM, thus fulfilling one of AMT’s stated aims: the assessment of biogeochemical models. Whilst the four datasets show broadly the same seasonal patterns and that the DCM shallows when surface chlorophyll increases, the depth and peak concentration of the DCM differ among datasets. For most of the datasets the column-integrated chlorophyll for both gyres is around 19 mg m-2 (with the AMT fluorescence-derived values being much lower); however the MEDUSA model has a disparity between the northern and southern gyres that is not understood. Although the seasonal increase in surface chlorophyll is tied to a commensurate decrease in concentration at depth, on an interannual basis years with enhanced surface levels of chlorophyll correspond to increases at depth. Satellite-derived observations of surface chlorophyll concentration act as a good predictor of interannual changes in DCM depth for both gyres during their autumn season, but provide less skill in spring. [ABSTRACT FROM AUTHOR]

Blooms of the red, filamentous cyanobacterium Planktothrix rubescens occur frequently in pre-alpine lakes in Europe, often with concomitant toxic microcystin (MC) production. Trophic transfer of MCs has been observed in bivalves, fish, and zooplankton species, while uptake of MCs into Diptera species could facilitate distribution of MCs into terrestrial food webs and habitats. In this study, we characterized a Planktothrix bloom in summer 2019 in Lake Mindelsee and tracked possible trophic transfer and/or bioaccumulation of MCs via analysis of phytoplankton, zooplankton (Daphnia) and emergent aquatic insects (Chaoborus, Chironomidae and Trichoptera). Using 16 S rRNA gene amplicon sequencing, we found that five sequence variants of Planktothrix spp. were responsible for bloom formation in September and October of 2019, and these MC-producing variants, provisionally identified as P. isothrix and/or P. serta, occurred exclusively in Lake Mindelsee (Germany), while other variants were also detected in nearby Lake Constance. The remaining cyanobacterial community was dominated by Cyanobiaceae species with high species overlap with Lake Constance, suggesting a well-established exchange of cyanobacteria species between the adjacent lakes. With targeted LC–HRMS/MS we identified two MC-congeners, MC-LR and [Asp3]MC-RR with maximum concentrations of 45 ng [Asp3]MC-RR/L in lake water in September. Both MC congeners displayed different predominance patterns, suggesting that two different MC-producing species occurred in a time-dependent manner, whereby [Asp3]MC-RR was clearly associated with the Planktothrix spp. bloom. We demonstrate an exclusive transfer of MC-LR, but not [Asp3]MC-RR, from phytoplankton into zooplankton reaching a 10-fold bioconcentration, yet complete absence of these MC congeners or their conjugates in aquatic insects. The latter demonstrated a limited trophic transfer of MCs from zooplankton to zooplanktivorous insect larvae (e.g., Chaoborus), or direct transfer into other aquatic insects (e.g. Chironomidae and Trichoptera), whether due to avoidance or limited uptake and/or rapid excretion of MCs by higher trophic emergent aquatic insects.

Satellite observations have given us a clear idea of the changes in chlorophyll in the surface ocean on both a seasonal and interannual basis, but repeated observations at depth are much rarer. The permanently-stratified subtropical gyres in the Atlantic are highly oligotrophic, with most production centred on a deep chlorophyll maximum (DCM) just above the nitracline. This study explores the variations in this feature in the core of both gyres, considering both seasonal and interannual variations, and the linkages between changes at the surface and sub-surface. The in situ observations come from the Atlantic Meridional Transect (AMT), a long-running UK monitoring programme, and also from biogeochemical Argo floats. AMT provides measurements spanning more than 25 years directed through the centres of these gyres, but samples only 2 to 4 months per year and thus cannot resolve the seasonal variations, whereas the profiling floats give coverage throughout the year, but without the rigid spatial repeatability. These observational records are contrasted with representation of the centres of the gyres in two different biogeochemical models: MEDUSA and ERSEM, thus fulfilling one of AMT’s stated aims: the assessment of biogeochemical models. Whilst the four datasets show broadly the same seasonal patterns and that the DCM shallows when surface chlorophyll increases, the depth and peak concentration of the DCM differ among datasets. For most of the datasets the column-integrated chlorophyll for both gyres is around 19 mg m-2 (with the AMT fluorescence-derived values being much lower); however the MEDUSA model has a disparity between the northern and southern gyres that is not understood. Although the seasonal increase in surface chlorophyll is tied to a commensurate decrease in concentration at depth, on an interannual basis years with enhanced surface levels of chlorophyll correspond to increases at depth. Satellite-derived observations of surface chlorophyll concentration act as a good predictor of interannual changes in DCM depth for both gyres during their autumn season, but provide less skill in spring.

Deep chlorophyll maxima (DCM) are a common phenomenon in low-trophic stratified lakes, and there is much less information about the occurrence of DCM in eutrophic and dystrophic lakes. Therefore, we performed in situ continuous measurements of chlorophyll concentration by a submersible spectrofluorometer in 23 temperate lakes with different trophic conditions (mesotrophic, eutrophic, and dystrophic). We perform chemical analyses of available nutrient forms to better understand the mechanism of DCM formation. We found the highest concentration of phytoplankton in the metalimnion or upper hypolimnion of most studied lakes, regardless of trophic conditions. Nevertheless, the differences in the number of phytoplankton between DCM and epilimnion were largest in low-trophic lakes, where the chl a concentration in DCM was even 15 times higher than that in the epilimnion. The mesotrophic lakes showed vertical niche segregation by phytoplankton, with a general pattern − green algae in the upper layer, below diatoms, then cryptophytes (and cyanobacteria). Our results indicated that the main factor for DCM in mesotrophic and eutrophic lakes was epilimnetic nutrient depletion, while in dystrophic lakes, it seems to be caused by other factors due to the depletion of dissolved nutrients in whole water profiles.

Deep-sea bamboo corals (family Isididae) have been used as archives for reconstructing changes in the past ocean. However, uncertainties remain regarding the interpretation of geochemical signals from their organic nodes, specifically the water depth of the signals recorded by the coral. Here we explore this question by measuring radiocarbon (14C) and nitrogen (15N) isotopic compositions of the organic nodes in six bamboo corals collected from the central and eastern tropical Atlantic between 700 m and 2000 m water depth. By comparing coral 14C to measured seawater data, regional shallow-water coral records and climate-model outputs, we find contrasting results between the two regions. Our bamboo coral 14C results from the eastern tropical Atlantic support previous studies that suggest organic node carbon is sourced primarily from the mixed layer of the ocean. By contrast, the 14C of bamboo coral organic nodes from the oligotrophic central Atlantic better correlates with the 14C content of the subsurface deep chlorophyll maximum layer rather than the surface mixed layer. Combined with nitrogen isotope data, this observation suggests that sinking and/or ambient zooplankton feeding on phytoplankton from the deep chlorophyll maximum layer can contribute a significant proportion of the diet of bamboo corals. These results suggest that the carbon source for bamboo corals organic nodes may not always reside in the mixed layer, especially in oligotrophic regions, which has implications for 14C-based age model development in bamboo corals. [ABSTRACT FROM AUTHOR]

Subsurface chlorophyll maximum layers (SCMLs) are nearly ubiquitous in stratified water columns and exist at horizontal scales ranging from the submesoscale to the extent of oligotrophic gyres. These layers of heightened chlorophyll and/or phytoplankton concentrations are generally thought to be a consequence of a balance between light energy from above and a limiting nutrient flux from below, typically nitrate (NO3). Here we present multiple lines of evidence demonstrating that iron (Fe) limits or with light colimits phytoplankton communities in SCMLs along a primary productivity gradient from coastal to oligotrophic offshore waters in the southern California Current ecosystem. SCML phytoplankton responded markedly to added Fe or Fe/light in experimental incubations and transcripts of diatom and picoeukaryote Fe stress genes were strikingly abundant in SCML metatranscriptomes. Using a biogeochemical proxy with data from a 40-y time series, we find that diatoms growing in California Current SCMLs are persistently Fe deficient during the spring and summer growing season. We also find that the spatial extent of Fe deficiency within California Current SCMLs has significantly increased over the last 25 y in line with a regional climate index. Finally, we show that diatom Fe deficiency may be common in the subsurface of major upwelling zones worldwide. Our results have important implications for our understanding of the biogeochemical consequences of marine SCML formation and maintenance.

The deep chlorophyll maximum (DCM) harbours diverse microbial communities and regulates carbon fixation and sequestration in marine ecosystems. Although variations in the DCM microbial community diversity and composition across oceans are reported, we still have a limited knowledge about the ecological mechanism driving the biogeography of DCM microbial communities and their contributions to ecosystem functioning.Here, based on DNA‐sequencing data (16S rRNA and 18S rRNA genes) with two size fractions collected from coastal to offshore regions of the South China sea (SCS), we found that both size‐fractionated bacterial and protistan communities in the DCM exhibited a distance‐decay pattern, which was strongly governed by deterministic processes (62.7%–72.2%) and mainly explained by environmental factors (34.1%–37.7%).Under warmer temperature and fresher salinity levels, the decreased abundance of protistan parasites induced the weakening parasitic interactions with other protists, whereas the increased abundance of protistan consumers enhanced the phagocytic interactions with bacteria. Such reconstruction of microbial communities induced by environmental variations was thought to constrain particulate organic carbon (POC) flux, as revealed by the partial least square regression model illustrating a strong correlation between the composition of functional protistan groups and POC content of DCM (R2 = 0.48; r = 0.75; p < 10−6).This work highlights the importance of environmental filtering in shaping DCM microbial communities and extends our understanding of the declined carbon flux of the DCM in the SCS under future scenarios. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

The Northwest Pacific Ocean (NWPO) is a significant sink for atmospheric CO2 but a paucity of large-scale phytoplankton surveys in the upper and lower euphotic zone results in uncertainties in estimates of the efficiency of the biological carbon pump there. Here, we report the spatial distribution of lipid biomarkers from diatoms (brassicasterol/epi-brassicasterol), dinoflagellates (dinosterol), and haptophytes (C37 alkenones) as proxies of phytoplankton biomass and community structure in suspended particles from the surface and deep chlorophyll maximum (DCM) layers across low- and mid-latitude regions of the NWPO. Our observations suggest that these lipid biomarkers can be used as indicators of the vertical distributions of phytoplankton biomass, which was comparable between the surface and DCM layers. Water masses with different nutrient concentrations strongly controlled the variations of lipid biomarkers, showing high biomass and the dominance of diatoms in the eutrophic Oyashio region, whereas low biomass and high proportions of dinoflagellates and haptophytes occurred in the oligotrophic Kuroshio region. Diatoms predominated in the DCM, likely enhancing carbon sequestration in the deep ocean and in sediments. Our results quantitatively demonstrate the horizontal and vertical variations of phytoplankton biomass and community structure, leading to an improved understanding of ecosystem function and biogeochemical cycles in this important region of the NWPO.

We examined the biogeochemical impact of paired mesoscale cyclones and anticyclones in spatial proximity (<200 km apart) in the North Pacific Subtropical Gyre. While previous studies have demonstrated that upwelling associated with the intensification of cyclonic eddies can supply nutrients supporting plankton productivity, we observed that steeper vertical gradients in inorganic nutrients increased nutrient fluxes due to diapycnal mixing during the mature stage of cyclonic eddies. The increased diapycnal nutrient supply was linked with expansion of eukaryotic phytoplankton biomass and intensification of the deep chlorophyll maximum (DCM) layer. This perturbation in the plankton community was associated with increased fluxes of biominerals (specifically particulate inorganic carbon and particulate silica) and isotopically enriched organic nitrogen in particles exported in the cyclone. The time‐integrated effects of thermocline vertical displacements on the lower euphotic zone were predictable deficits and surpluses of inorganic nutrients and dissolved oxygen, respectively. However, the stoichiometry of oxygen and inorganic nutrients differed from that predicted for production and consumption of phytoplankton biomass, requiring additional biological processes that decouple changes in oxygen and nutrient concentrations. The dynamics revealed by this study may be a common feature of oligotrophic ecosystems, where mesoscale biogeochemical perturbations are buffered by the DCM layer, which limits the ecological impact of eddies in the well‐lit, near‐surface ocean. Key Points: The steepness of the nutricline in mature mesoscale eddies is an important driver of variability in pelagic ecosystemsDifferences in eukaryotic phytoplankton communities alter the export of particulate inorganic carbon and particulate silica across eddiesRelative changes in oxygen and nutrients linked to displacements of the thermocline reflect anomalous cycling of organic matter [ABSTRACT FROM AUTHOR]

Context: Ecotones are boundary zones formed where overlap between neighboring ecosystems creates an intermediate ecosystem with unique ecological characteristics. Dynamic ecotones change position along a boundary over time and can be further categorized as either shifting, where the adjacent ecosystems alternatively drive movement of the ecotone but maintain the same relative location over time, or directional, where one system encroaches into the other and the ecotone moves laterally. Objectives: The purpose of this work was to examine how climate change alters movement dynamics of both directional and shifting ecotones. Methods: In three ecosystem case studies, we examine the effects of climate change on landscape-scale ecotone movement across the marine, terrestrial, and interfacing environments. Results: Shifts in local and global climate drive changes in ecotone patterns, increasing directional ecotone movement at both shifting and directional ecotones. Specifically, unidirectional changes in climate patterns disrupt dynamic equilibria at shifting ecosystem boundaries, thereby facilitating unidirectional movement at the previously shifting boundaries. Climate changes additionally accelerate pre-existing directional migration of ecotones through changes to abiotic gradients. Conclusion: Directional climate change increases directional movement in multiple types of ecotone. Future work should consider the rate and feedback mechanisms of ecotone movement and function at additional ecotones. [ABSTRACT FROM AUTHOR]

Planktothrix rubescens is a harmful planktonic cyanobacterium, forming concentrated metalimnetic populations in deep oligo- and mesotrophic lakes, even after successful restoration. In Lake Zurich (Switzerland), P. rubescens emerged as a keystone species with annual mass developments since the 1970s. Its success was partly attributed to effects of lake warming, such as changes in thermal stratification and seasonal deep mixing. However, recent observations based on a biweekly monitoring campaign (2009–2020) revealed two massive breakdowns and striking seasonal oscillations of the population. Here, we disentangle positive from negative consequences of secular lake warming and annual variations in weather conditions on P. rubescens dynamics: (i) despite the high survival rates of overwintering populations (up to 25%) during three consecutive winters (2014–2016) of incomplete deep convective mixing, cyanobacterial regrowth during the following stratified season was moderate and not overshooting a distinct standing stock threshold. Moreover, we recorded a negative trend for annual population maxima and total population size, pointing to a potential nutrient limitation after a series of incomplete winter mixing. Thus, the predication of steadily increasing blooms of P. rubescens could not be confirmed for the last decade. (ii) The seasonal reestablishment of P. rubescens was strongly coupled with a timely formation of a stable metalimnion structure, where the first positive net growth in the following productive summer season was observed. The trigger for the vertical positioning of filaments within the metalimnion was irradiance and not maximal water column stability. Repetitive disruptions of the vernal metalimnion owing to unstable weather conditions, as in spring 2019, went in parallel with a massive breakdown of the standing stock and marginal regrowth during thermal stratification. (iii) Driven by light intensity, P. rubescens was entrained into the turbulent epilimnion in autumn, followed by a second peak in population growth. Thus, the typical bimodal growth pattern was still intact during the last decade. Our long-term study highlights the finely tuned interplay between climate-induced changes and variability of thermal stratification dynamics and physiological traits of P. rubescens , determining its survival in a mesotrophic temperate lake. [ABSTRACT FROM AUTHOR]

Deep Chlorophyll Maxima (DCM) are ubiquitous features in stratified oceanic systems. Their establishment and maintenance result from hydrographical stability favoring specific environmental conditions with respect to light and nutrient availability required for phytoplankton growth. This stability can potentially be challenged by mesoscale eddies impacting the water column's vertical structure and thus the environmental parameters that condition the subsistence of DCMs. Here, data from the global BGC‐Argo float network are collocated with mesoscale eddies to explore their impact on DCMs. We show that cyclonic eddies, by providing optimal light and nutrient conditions, increase the occurrence of DCMs characterized by Deep Biomass Maxima for phytoplankton. In contrast, DCMs in anticyclonic eddies seem to be driven by photoacclimation as they coincide with Deep Acclimation Maxima without biomass accumulation. These findings suggest that the two types of eddies potentially have different impacts on the role of DCMs in global primary production. Plain Language Summary: In the global ocean, phytoplankton can be found at depths where their growth is limited by their access to light from the surface and nutrient supply from below. Depending on the combination of environmental conditions, phytoplankton at depth can either accumulate in densely populated layers or deploy adaptive survival strategies such as multiplying their internal light sensors. Given their dependence on environmental variations, phytoplankton can be affected by physical disturbances that alter the vertical structure of the water column's properties. Among these are mesoscale eddies, large water structures dominated by rotation on the horizontal plane which also induce vertical physical movements. In this study, we associate data from autonomous robots that drift with ocean currents and sample physical and biological properties down to a depth of 2,000 meters, with mesoscale eddies detected by satellite. We show that on a global scale, cold mesoscale eddies promote the growth of phytoplankton at depth. Key Points: Deep Chlorophyll Maxima are key biological features of stratified environments that escape satellite ocean‐color detectionThe impact of mesoscale eddies on DCM is assessed by matching a global BGC‐Argo database with eddies detected from satellite altimetryWhile cyclonic eddies increase biomass maxima occurrences, anticyclonic eddies favor the presence of maxima resulting from photoacclimation [ABSTRACT FROM AUTHOR]

The Cooperative Institute for Great Lakes Research (CIGLR) in collaboration with the Great Lakes Observing System and National Oceanic and Atmospheric Administration, Great Lakes Environmental Research Laboratory (NOAA GLERL) deployed an autonomous underwater glider in southern Lake Michigan several times per year between 2012 and 2019 to collect offshore (>30 m depth) limnological measurements, including temperature, photosynthetically active radiation (beginning during 2015), and chlorophyll -α fluorescence. From these data, we calculated mixed layer depth, several measures of light penetration (diffuse attenuation coefficient, first optical depth, euphotic zone depth), and depth of the subsurface chlorophyll -α maxima. During summer, mean offshore mixed layer depth was typically 10–15 m, K d for PAR was 0.1–0.17 m−1, first optical depth was 6–9 m, euphotic zone depth was 35–40 m, and depth of subsurface chlorophyll -α maxima was 30–35 m. We also observed substantial spatial and temporal variation in these values across the basin and within and among seasons. Glider-based observations provide a wider horizontal and vertical perspective than other methods (e.g., ship- and satellite-based observations, buoys, and fixed moorings), and are therefore a valuable, complementary tool for Great Lakes limnology. The set of observations reported here provide seasonal and basin-scale information that may help to identify anomalies useful for future glider-assisted investigation into the role of biophysical processes in Great Lakes limnology and ecology. [ABSTRACT FROM AUTHOR]

Within the framework of the Marine Ecosystem Assessment for the Southern Ocean (MEASO), this paper brings together analyses of recent trends in phytoplankton biomass, primary production and irradiance at the base of the mixed layer in the Southern Ocean and summarises future projections. Satellite observations suggest that phytoplankton biomass in the mixed-layer has increased over the last 20 years in most (but not all) parts of the Southern Ocean, whereas primary production at the base of the mixed-layer has likely decreased over the same period. Different satellite models of primary production (Vertically Generalised versus Carbon Based Production Models) give different patterns and directions of recent change in net primary production (NPP). At present, the satellite record is not long enough to distinguish between trends and climate-related cycles in primary production. Over the next 100 years, Earth system models project increasing NPP in the water column in the MEASO northern and Antarctic zones but decreases in the Subantarctic zone. Low confidence in these projections arises from: (1) the difficulty in mapping supply mechanisms for key nutrients (silicate, iron); and (2) understanding the effects of multiple stressors (including irradiance, nutrients, temperature, pCO2, pH, grazing) on different species of Antarctic phytoplankton. Notwithstanding these uncertainties, there are likely to be changes to the seasonal patterns of production and the microbial community present over the next 50–100 years and these changes will have ecological consequences across Southern Ocean food-webs, especially on key species such as Antarctic krill and silverfish.

Accurate estimation of the underwater light field associated with photosynthetically available radiation (PAR) is critical to compute phytoplankton growth rate and net primary production (NPP), and to assess photo‐physiological response of phytoplankton, such as changes in cellular pigmentation. However, methods to estimate PAR used in many previous studies lack in accuracy, likely resulting in significant bias in light‐dependent products such as NPP derived from remote sensing, model simulations, or autonomous platforms. Here we propose and validate a new model for more accurate estimation of the subsurface PAR profile which uses chlorophyll concentration as its input. Validation is performed using 1,744 BGC‐Argo profiles of chlorophyll fluorescence that are calibrated with surface satellite‐derived chlorophyll concentration over their lifetime. The independent verification with the float's PAR sensors confirms the accuracy of satellite chlorophyll estimate worldwide and in the Southern Ocean in particular. Plain Language Summary: In this study, we propose a new model to compute the underwater light field available for phytoplankton growth. The model uses as its inputs the above water radiation (which can be obtained from public databases) and the subsurface distribution of chlorophyll‐a, a pigment shared by all phytoplankton that can be estimated from sensors deployed on robotic platforms and that is used by many ocean ecosystem models. We test the model accuracy using data observed by the light sensors on autonomous profiling floats and find it performs well. The comparison also highlights that estimates of surface chlorophyll concentrations from satellites are unbiased worldwide, in contrast to some published accounts. Key Points: The model takes into account the vertical change of chlorophyll and improves the estimate of the subsurface photosynthetically available radiation fieldThe model is applicable to autonomous platforms, ecosystem and biogeochemical models, and ocean color remote sensingModel comparison with independent in‐situ radiometry confirms that satellite chlorophyll‐a retrievals are unbiased worldwide [ABSTRACT FROM AUTHOR]

Stratified oceanic systems are characterized by the presence of a so‐called Deep Chlorophyll a Maximum (DCM) not detectable by ocean color satellites. A DCM can either be a phytoplankton (carbon) biomass maximum (Deep Biomass Maximum, DBM), or the consequence of photoacclimation processes (Deep photoAcclimation Maximum, DAM) resulting in the increase of chlorophyll a per phytoplankton carbon. Even though these DCM (further qualified as either DBMs or DAMs) have long been studied, no global‐scale assessment has yet been undertaken and large knowledge gaps still remain in relation to the environmental drivers responsible for their formation and maintenance. In order to investigate their spatial and temporal variability in the open ocean, we use a global data set acquired by more than 500 Biogeochemical‐Argo floats given that DCMs can be detected from the comparative vertical distribution of chlorophyll a concentrations and particulate backscattering coefficients. Our findings show that the seasonal dynamics of the DCMs are clearly region‐dependent. High‐latitude environments are characterized by a low occurrence of intense DBMs, restricted to summer. Meanwhile, oligotrophic regions host permanent DAMs, occasionally replaced by DBMs in summer, while subequatorial waters are characterized by permanent DBMs benefiting from favorable conditions in terms of both light and nutrients. Overall, the appearance and depth of DCMs are primarily driven by light attenuation in the upper layer. Our present assessment of DCM occurrence and of environmental conditions prevailing in their development lay the basis for a better understanding and quantification of their role in carbon budgets (primary production and export). Key Points: The main characteristics and drivers of Deep Chlorophyll Maxima (DCM) are analyzed from a global BGC‐Argo databaseLatitude and season determine the occurrence and characteristics of DCMsDCMs result from photoacclimation or biomass accumulation, depending on the availability of light and nitrate [ABSTRACT FROM AUTHOR]

Blooms of the red, filamentous cyanobacterium Planktothrix rubescens occur frequently in pre-alpine lakes in Europe, often with concomitant toxic microcystin (MC) production. Trophic transfer of MCs has been observed in bivalves, fish, and zooplankton species, while uptake of MCs into Diptera species could facilitate distribution of MCs into terrestrial food webs and habitats. In this study, we characterized a Planktothrix bloom in summer 2019 in Lake Mindelsee and tracked possible trophic transfer and/or bioaccumulation of MCs via analysis of phytoplankton, zooplankton (Daphnia) and emergent aquatic insects (Chaoborus , Chironomidae and Trichoptera). Using 16 S rRNA gene amplicon sequencing, we found that five sequence variants of Planktothrix spp. were responsible for bloom formation in September and October of 2019, and these MC-producing variants, provisionally identified as P. isothrix and/or P. serta , occurred exclusively in Lake Mindelsee (Germany), while other variants were also detected in nearby Lake Constance. The remaining cyanobacterial community was dominated by Cyanobiaceae species with high species overlap with Lake Constance, suggesting a well-established exchange of cyanobacteria species between the adjacent lakes. With targeted LC–HRMS/MS we identified two MC-congeners, MC-LR and [Asp3]MC-RR with maximum concentrations of 45 ng [Asp3]MC-RR/L in lake water in September. Both MC congeners displayed different predominance patterns, suggesting that two different MC-producing species occurred in a time-dependent manner, whereby [Asp3]MC-RR was clearly associated with the Planktothrix spp. bloom. We demonstrate an exclusive transfer of MC-LR, but not [Asp3]MC-RR, from phytoplankton into zooplankton reaching a 10-fold bioconcentration, yet complete absence of these MC congeners or their conjugates in aquatic insects. The latter demonstrated a limited trophic transfer of MCs from zooplankton to zooplanktivorous insect larvae (e.g., Chaoborus), or direct transfer into other aquatic insects (e.g. Chironomidae and Trichoptera) , whether due to avoidance or limited uptake and/or rapid excretion of MCs by higher trophic emergent aquatic insects. [Display omitted] • A Planktothrix sp. bloom specific to Lake Mindelsee produced the toxin [Asp3]MC-RR. • MC-LR-transfer from Microcystis sp. to zooplankton resulted in MC-biomagnification. • MC or MC metabolite-trophic transfer to emergent aquatic insects was not detected. • No toxic Planktothrix sp. blooms were reported in the adjacent Lake Constance. • Synechococcus phylotypes of Lake Mindelsee are potentially exchanged between lakes. [ABSTRACT FROM AUTHOR]

Quantification of the vertical distribution of chlorophyll a (Chl- a), as one of characteristics of primary production (PP), is critically important to estimate annual PP in the water column (IPP) using models and remote sensing data. IPP estimation in optically complex and highly variable waters such as the Kara Sea is not a trivial task. In the present study, based on the data obtained during three multidisciplinary cruises to the eastern regions of the Kara Sea in August–October, the differences in the vertical distribution of Chl- a and PP under and without the influence of the river plume (Case II and Case I water types, respectively) are established. In Case I waters in August 2014, under the low values of the diffuse attenuation coefficient (K d) of downwelling photosynthetically available radiation (PAR) (the median value (Me) K d = 0.158 m-1) and the deep euphotic layer (Z eu) (Me = 30 m), the deep chlorophyll maximum (DCM) was well-pronounced. At the end of September 2015 in Case II waters influenced by the river runoff, when Me K d increased 1.7-fold and Z eu decreased 1.3-fold, the DCM was absent. Also, the DCM was not manifested at the end of the growing season, under conditions of extremely low underwater PAR (Me = 0.35 mol quanta m−2 d−1). In the sampling period, the PP maxima were observed at the surface and the DCM did not influence the vertical PP distribution. The depth of the nitracline was directly associated with the distribution of riverine waters and determined the depth and degree of DCM manifestation. The outcomes of the presented study suggest that a decrease in subsurface PAR influenced by the impact of riverine waters and the total decline of incident radiation from August to October determine the vertical distribution of PP and Chl- a. • The vertical distribution of PP and Chl- a depended on the river plume dynamics. • DCM was not observed in the freshwater runoff period. • A decrease in PAR at the end of the growing season led to DCM dissipation. • Underwater PAR was the major abiotic factor influencing vertical distribution. [ABSTRACT FROM AUTHOR]

Bio-optical data, obtained during six cruises in the Black Sea carried out during periods of seasonal stratification in years between 1996 and 2016, have been used to parametrize phytoplankton light absorption ($${a_{ph}}\left({\rm{\lambda }} \right)$$) in the deep chlorophyll maximum (DCM) layer located near the bottom of euphotic zone. Relationships between $${a_{ph}}\left({\rm{\lambda }} \right)$$ and the sum of chlorophyll-a and phaeopigment concentrations (Chl-a) differed from those for the summertime upper mixed layer (UML). Notably, chlorophyll a specific absorption coefficients ($$a_{ph}^\ast\left(\lambda \right)$$) were lower in the DCM and more comparable with $$a_{ph}^\ast\left(\lambda \right)$$ values typical for winter phytoplankton in the Black Sea. The $${a_{ph}}\left({\rm{\lambda }} \right)$$ spectral shapes in the DCM differed markedly from those in winter and in the summer UML, due to a shoulder at ~490 nm and a local maximum at ~550 nm corresponding to the absorption bands of phycourobilin and phycoerythrobilin. Light absorbing properties of phytoplankton in the DCM (amplitude and spectral shape of $$a_{ph}^\ast\left(\lambda \right)$$) reflected physiological acclimation to local conditions on the cellular level and population shifts leading to changes in the biomass-dominant species, with Synechococcus spp. domination in the DCM. The parameterization of phytoplankton absorption in the DCM will enable refined spectral models of the downwelling radiance and primary production in the Black Sea.

During the late Miocene the Mediterranean experienced a dramatic intensification of opaline accumulation, recorded by the deposition of diatomaceous sediments. The fine lamination of these deposits potentially records annual to sub‐annual palaeoceanographic processes that occurred during a critical phase of the geodynamic evolution of the Mediterranean basin, which eventually led to the Messinian salinity crisis. The diatomaceous facies has been interpreted by previous researchers as the product of intensified upwelling currents and of bottom anoxia formation in the Mediterranean basin. However, until now, no efforts have been made to unravel the sedimentological and micropalaeontological content of these deposits at the lamina‐scale. This paper presents the first case study of a systematic scanning electron microscope‐based morphological investigation of the diatomaceous sediments deposited during the late Miocene at the northernmost offshoot of the Mediterranean basin (Piedmont Basin, north‐west Italy). Using a non‐invasive analytical approach, six faciological components (laminae, laminated packets, non‐laminated intervals, burrows, opal‐rich aggregates and mixed pelletal structures) and their relationships are described and interpreted herein. Following the lamina‐scale study of these sediments, an annual sedimentary cycle could be identified and an accumulation rate (ca 50 cm kyr−1) inferred that is atypical for a setting actively influenced by upwelling. The role played by the entanglement of diatom valves in creating a physical barrier to the bioturbation is here emphasized as the main process responsible for the preservation of the laminated fabric of diatomaceous sediments, challenging the supposed role of deep anoxia. These results suggest that the late Miocene diatomaceous deposition in the Piedmont Basin cannot be univocally considered as a by‐product of upwelling intensification and seafloor oxygen depletion. [ABSTRACT FROM AUTHOR]

A deep chlorophyll layer (DCL) is a common feature of many deep, oligotrophic lakes including Lake Superior. Mechanisms generating and maintaining DCLs are variable across lakes, and seasonal patterns and relationships of DCL structure to physical variables are not well described. Using vertical profile data for physical and biological variables from western and central Lake Superior, we described seasonal patterns in DCL structure and other physical and biological parameters and applied linear mixed-effects models to determine how different physical factors (surface temperature, thermocline depth, and 1% photosynthetically active radiation (PAR) depth) affect the depth, thickness, maximum concentration, and integrated chlorophyll of the DCL. We observed clear seasonal patterns in the development and degradation of the DCL that coincide with seasonal changes in light and temperature. Modeling analysis using linear mixed-effects models showed that the DCL thickness was best predicted by surface temperature (R2 = 0.51) followed by thermocline depth (R2 = 0.36), and the deep chlorophyll maximum (DCM) concentration was best predicted by surface temperature (R2 = 0.26). The 1% PAR depth was not implicated as an important predictor, but observations from seasonal data suggest that it plays a role in the depth of the DCM. While no relationship was found between surface temperature and DCL-integrated chlorophyll, DCL thickness decreased and DCM concentration increased with increasing surface temperature, which could have implications for productivity in the DCL as the lake continues to warm. [ABSTRACT FROM AUTHOR]

Mesoscale eddies can have a strong impact on regional biogeochemistry and primary productivity. To investigate the effect of the upwelling of seawater by western Pacific eddies on the composition of the active planktonic marine archaeal community composition of the deep chlorophyll maximum (DCM) layer, mesoscale cold-core eddies were simulated in situ by mixing western Pacific DCM layer water with mesopelagic layer (400 m) water. Illumina sequencing of the 16S rRNA gene and 16S rRNA transcripts indicated that the specific heterotrophic Marine Group IIb (MGIIb) taxonomic group of the DCM layer was rapidly stimulated after receiving fresh substrate from 400 m water, which was dominated by uncultured autotrophic Marine Group I (MGI) archaea. Furthermore, niche differentiation of autotrophic ammonia-oxidizing archaea (MGI) was demonstrated by deep sequencing of 16S rRNA, amoA, and accA genes, respectively. Similar distribution patterns of active Marine Group III (MGIII) were observed in the DCM layer with or without vertical mixing, indicating that they are inclined to utilize the substrates already present in the DCM layer. These findings underscore the importance of mesoscale cyclonic eddies in stimulating microbial processes involved in the regional carbon cycle.

Numerical modeling is a key tool to complement the current physical and biogeochemical observational datasets. It is essential for understanding the role of oceanographic processes on marine food webs and producing climate change projections of variables affecting key ecosystem functions. In this work, we evaluate the horizontal and vertical patterns of four state-of-the-art coupled physical–biogeochemical models, three of them already published. Two of the models include data assimilation, physical and/or biological, and two do not. Simulations are compared to the most exhaustive dataset of in situ observations in the North Western Mediterranean, built ad hoc for this work, comprising gliders and conventional CTD surveys and complemented with satellite observations. The analyses are performed both in the whole domain and in four subregions (Catalan Shelf, Ebro Delta, Mallorca Channel, and Ibiza Channel), characterized by a priori divergent primary production dynamics and driving mechanisms. Overall, existing models offer a reasonable representation of physical processes including stratification, surface temperature, and surface currents, but it is shown that relatively small differences among them can lead to large differences in the response of biogeochemical variables. Our results show that all models are able to reproduce the main seasonal patterns of primary production both at the upper layer and at the deep chlorophyll maximum (DCM), as well as the differential behavior of the four subregions. However, there are significant discrepancies in the local variability of the intensity of the winter mixing, phytoplankton blooms, or the intensity and depth of the DCM. All model runs show markedly contrasting patterns of interannual phytoplankton biomass in all four subregions. This lack of robustness should dissuade end users from using them to fill gaps in time series observations without assessing their appropriateness. Finally, we discuss the usability of these models for different applications in marine ecology, including fishery oceanography.

Abstract Background The photic zone of aquatic habitats is subjected to strong physicochemical gradients. To analyze the fine-scale variations in the marine microbiome, we collected seven samples from a single offshore location in the Mediterranean at 15 m depth intervals during a period of strong stratification, as well as two more samples during the winter when the photic water column was mixed. We were able to recover 94 new metagenome-assembled genomes (MAGs) from these metagenomes and examine the distribution of key marine microbes within the photic zone using metagenomic recruitment. Results Our results showed significant differences in the microbial composition of different layers within the stratified photic water column. The majority of microorganisms were confined to discreet horizontal layers of no more than 30 m (stenobathic). Only a few such as members of the SAR11 clade appeared at all depths (eurybathic). During the winter mixing period, only some groups of bloomers such as Pseudomonas were favored. Although most microbes appeared in both seasons, some groups like the SAR116 clade and some Bacteroidetes and Verrucomicrobia seemed to disappear during the mixing period. Furthermore, we found that some microbes previously considered seasonal (e.g., Archaea or Actinobacteria) were living in deeper layers within the photic zone during the stratification period. A strong depth-related specialization was detected, not only at the taxonomic level but also at the functional level, even within the different clades, for the manipulation and uptake of specific polysaccharides. Rhodopsin sequences (green or blue) also showed narrow depth distributions that correlated with the taxonomy of the microbe in which they were found but not with depth. Conclusions Although limited to a single location in the Mediterranean, this study has profound implications for our understanding of how marine microbial communities vary with depth within the photic zone when stratified. Our results highlight the importance of collecting samples at different depths in the water column when comparing seasonal variations and have important ramifications for global marine studies that most often take samples from only one single depth. Furthermore, our perspective and approaches (metagenomic assembly and recruitment) are broadly applicable to other metagenomic studies.

Mesoscale eddies can have a strong impact on regional biogeochemistry and primary productivity. To investigate the effect of the upwelling of seawater by western Pacific eddies on the composition of the active planktonic marine archaeal community composition of the deep chlorophyll maximum (DCM) layer, mesoscale cold-core eddies were simulated in situ by mixing western Pacific DCM layer water with mesopelagic layer (400 m) water. Illumina sequencing of the 16S rRNA gene and 16S rRNA transcripts indicated that the specific heterotrophic Marine Group IIb (MGIIb) taxonomic group of the DCM layer was rapidly stimulated after receiving fresh substrate from 400 m water, which was dominated by uncultured autotrophic Marine Group I (MGI) archaea. Furthermore, niche differentiation of autotrophic ammonia-oxidizing archaea (MGI) was demonstrated by deep sequencing of 16S rRNA, amoA , and accA genes, respectively. Similar distribution patterns of active Marine Group III (MGIII) were observed in the DCM layer with or without vertical mixing, indicating that they are inclined to utilize the substrates already present in the DCM layer. These findings underscore the importance of mesoscale cyclonic eddies in stimulating microbial processes involved in the regional carbon cycle. [ABSTRACT FROM AUTHOR]

Subsurface chlorophyll maxima (SCM) layers are becoming more important to the Arctic shelf ecosystem as phytoplankton growing season and ice‐free water increase. We measured size‐fractionated pigments and environmental variables on the western Arctic shelf during the open‐water season in 2014, to study the spatial pattern and mechanisms of the phytoplankton community at the SCM. The SCM layers accumulated approximately 50% of the chlorophyll a (Chl a) standing stock in the water column, and the contribution increased to >70% for high‐biomass areas, suggesting the biological importance of the summer SCM in the shelf ecosystem. The spatial difference in phytoplankton biomass was caused mainly by bioavailable nitrogen concentrations, and the spatial difference in diatom size may be the result of differences in predominant species. High biomass and large diatoms dominated in the SCM layers, making the south and north of the Chukchi shelf a biological hotspot for the ecosystem's food chain and benthos of the western Arctic shelf in summer. Plain Language Summary: Subsurface chlorophyll maxima (SCM) play an increasingly important role in shelf ecosystems in the western Arctic Ocean. The correlations between the subsurface phytoplankton community and environmental variables were examined to gain insight into the mechanisms underlying phytoplankton distribution in SCM layers. Spatial variability of the chlorophyll a (Chl a) standing stock showed a high phytoplankton biomass in the south Bering shelf (SBS), south Chukchi shelf (SCS), and north Chukchi shelf (NCS). Small diatoms dominated the phytoplankton community in the SBS, whereas large diatoms dominated in the SCS and NCS. The spatial difference in phytoplankton biomass was caused mainly by bioavailable nitrogen concentrations, and the spatial difference in diatom size may be the result of differences in predominant species. Summer SCM accumulate a large number of large diatoms in the water column, making the south and north of the Chukchi shelf a biological hotspot for the marine ecosystem on the western Arctic shelf in summer. Key Points: Summer SCM accumulate ~50% of the total water column chlorophyll aNitrate availability causes the spatial difference in phytoplankton biomassHigh biomass and dominance of large diatoms make the south and north of the Chukchi shelf a biological hotspot in summer [ABSTRACT FROM AUTHOR]

Chlorophyll measurements are commonly used to estimate phytoplankton biomass. However, phytoplankton readily acclimate to variations in light through a range of phenotypic responses, including major adjustments in chlorophyll pigmentation at the cellular level. The ratio of pigment chlorophyll to carbon concentration (Chl:C) is a commonly used metric in the oceanographic community to explore photoacclimation responses to varied light levels, yet is relatively rare in freshwater studies. Here we explore how nutrient variability impacted summertime Chl:C ratios of a natural phytoplankton community throughout the water column of a stratified oligotrophic lake. We utilized both long-term (18–24 years) and high-frequency (daily) data from Crater Lake, Oregon, a deep mountain lake with little anthropogenic disturbance. As expected, fluctuation in nutrients had a strong impact on phytoplankton particle density, primary productivity, light penetration, and water clarity. However, chlorophyll concentration did not register predictable changes even though the vertical location of the deep chlorophyll maximum was responsive to the overlying algal density. The impact of elevated nutrients on the Chl:C ratio was further complicated by upward shifts in chlorophyll distribution. The muted response of chlorophyll concentration to nutrients may be partially explained by variations in phytoplankton community composition or iron stress. [ABSTRACT FROM AUTHOR]

In the Indian Ocean regions under the influence of monsoons, two phytoplankton blooms characterize the seasonal cycle of surface chlorophyll, one during summer, and the other during winter. In the Northern Mozambique Channel, however, where the wind regime is an extension of the northern Indian Ocean monsoons, the annual cycle of chlorophyll concentrations is characterized by a single winter bloom. Wind stress and surface net heat flux modulate the seasonal cycle of the mixed layer depth with impacts on the surface chlorophyll. In order to evaluate the importance of these forcing fields on the seasonality of the mixed layer depth, and consequently, the surface chlorophyll variability, we used a suite of physical-biogeochemical model sensitivity experiments. Our results show that the seasonal cycle of surface chlorophyll is primarily modulated by the net heat flux while the wind field controls the amplitude. The winter bloom is triggered by negative surface heat fluxes, where cooling at the surface induces mixing and entrainment of nutrients at the base of the nutricline and light is not limiting. Winds enhance the winter bloom by uplifting additional nutrients and diluting subsurface chlorophyll into the surface layer. In the summertime, weaker wind stress and positive heat flux inhibit vertical mixing. As a consequence, the surface layer is depleted in nutrients and a deep chlorophyll maximum is formed. Analysis of top-down control on phytoplankton biomass reveals that zooplankton abundance increases in a near-linear proportion with phytoplankton biomass despite the deepening of the mixed layer depth. This suggests that the phytoplankton stock in the Northern Mozambique Channel is also controlled by grazing, given that zooplankton biomass is not directly affected by the deepening of the mixed layer during wintertime. [ABSTRACT FROM AUTHOR]

Changes in phytoplankton community composition and structure can have broad-scale ecosystem effects; however, drivers of species diversity in planktonic systems are not well understood. In lakes, a common but not thoroughly tested assumption is that shallow, nearshore waters are much more diverse and productive, and contribute considerably more material and energy to pelagic food webs than deeper waters farther offshore. Lake Superior is a large, cold, oligotrophic freshwater system which can provide insight into community organization under oligotrophic conditions. We used epilimnion and deep chlorophyll layer phytoplankton data from a lake-wide sampling program conducted in 2011 and 2016 to test whether assemblage composition, total algal biovolume, cell concentrations, diversity, and richness vary with depth. Although lake depth was an important factor in structuring assemblage composition, there were no clear nearshore-offshore gradients in cell density or biovolume despite the exposure of nearshore areas to higher concentrations of watershed-derived nutrients. Shannon diversity increased slightly with increasing depth, whereas richness was uncorrelated. Understanding of the nearshore-offshore patterns in phytoplankton community characteristics in the Great Lakes has implications for designing monitoring strategies and for considering how further changes in climate and nutrient deposition would affect the base of the food web. [ABSTRACT FROM AUTHOR]

Observing the vertical dynamic of phytoplankton in the water column is essential to understand the evolution of the ocean primary productivity under climate change and the efficiency of the CO2 biological pump. This is usually made through in-situ measurements. In this paper, we propose a machine learning methodology to infer the vertical distribution of phytoplankton pigments from surface satellite observations, allowing their global estimation with a high spatial and temporal resolution. After imputing missing values through iterative completion Self-Organizing Maps, smoothing and reducing the vertical distributions through principal component analysis, we used a Self-Organizing Map to cluster the reduced profiles with satellite observations. These referent vector clusters were then used to invert the vertical profiles of phytoplankton pigments. The methodology was trained and validated on the MAREDAT dataset and tested on the Tara Oceans dataset. The different regression coefficients R2 between observed and estimated vertical profiles of pigment concentration are, on average, greater than 0.7. We could expect to monitor the vertical distribution of phytoplankton types in the global ocean.

Abstract Dissolved oxygen is a key player in water quality. Stratified water bodies show distinct vertical patterns of oxygen concentration, which can originate from physical, chemical or biological processes. We observed a pronounced metalimnetic oxygen minimum in the low-nutrient Rappbode Reservoir, Germany. Contrary to the situation in the hypolimnion, measurements of lateral gradients excluded the sediment contact zone from the major sources of oxygen depletion for the metalimnetic oxygen minimum. Instead, the minimum was the result of locally enhanced oxygen consumption in the open water body. A follow-up monitoring included multiple chlorophyll a fluorescence sensors with high temporal and vertical resolution to detect and document the evolution of phytoplankton. While chlorophyll fluorescence sensors with multiple channels detected a mass development of the phycoerythrin-rich cyanobacterium Planktothrix rubescens in the metalimnion, this species was overlooked by the commonly used single-channel chlorophyll sensor. The survey indicated that the waning P. rubescens fluorescence was responsible for the oxygen minimum in the metalimnion. We hypothesize that pelagic processes, i.e., either oxygen use through decomposition of dead organic material originating from P. rubescens or P. rubescens extending its respiration beyond its photosynthetic activity, induced the metalimnetic oxygen minimum. The deeper understanding of the oxygen dynamics is mandatory for optimizing reservoir management. Graphical abstract Image 1 Highlights • A metalimnetic oxygen minimum (MOM) down to 40% saturation was observed in summer. • The MOM was not imported from the sediment at the side walls. • The MOM was a consequence of pelagic processes, such as respiration in the metalimnion. • We hypothesize that Planktothrix rubescens in the metalimnion caused the MOM. [ABSTRACT FROM AUTHOR]

Bio-optical data, obtained during six cruises in the Black Sea carried out during periods of seasonal stratification in years between 1996 and 2016, have been used to parametrize phytoplankton light absorption () in the deep chlorophyll maximum (DCM) layer located near the bottom of euphotic zone. Relationships between and the sum of chlorophyll-a and phaeopigment concentrations (Chl-a) differed from those for the summertime upper mixed layer (UML). Notably, chlorophyll a specific absorption coefficients () were lower in the DCM and more comparable with values typical for winter phytoplankton in the Black Sea. The spectral shapes in the DCM differed markedly from those in winter and in the summer UML, due to a shoulder at ~490 nm and a local maximum at ~550 nm corresponding to the absorption bands of phycourobilin and phycoerythrobilin. Light absorbing properties of phytoplankton in the DCM (amplitude and spectral shape of ) reflected physiological acclimation to local conditions on the cellular level and population shifts leading to changes in the biomass-dominant species, with Synechococcus spp. domination in the DCM. The parameterization of phytoplankton absorption in the DCM will enable refined spectral models of the downwelling radiance and primary production in the Black Sea. [ABSTRACT FROM AUTHOR]

Ecological traits of phytoplankton are being incorporated into models to better understand the dynamics of marine ecosystems and to predict their response to global change. We have compared the distribution of major phytoplankton groups in two different systems: in surface waters of the NW Mediterranean during key ecological periods, and in the DCM (deep chlorophyll maximum) formed in summer in the temperate NE Atlantic. This comparison disentangled the influence of light and nutrients on the relative position of diatoms, dinoflagellates, prymnesiophytes, pelagophytes, chlorophytes, Synechococcus and Prochlorococcus in these environments. Three clusters formed according to their affinity for nutrients: diatoms, chlorophytes and dinoflagellates as the most eutrophic groups; Synechococcus, pelagophytes and prymnesiophytes as mesotrophic groups; and Prochlorococcus as an oligotrophic group. In terms of irradiance, the phytoplankton groups did not cluster clearly. Comparing the nutrient and light preferences of the groups with their distribution in the DCM, dinoflagellates and chlorophytes appear as the most stressed, i.e. their position was most distant from their optimal light and nutrient conditions. Diatoms stayed in deeper than optimal irradiance layers, probably to meet their high nutrient requirements. On the opposite side, low nutrient requirements allowed Prochlorococcus to remain in the uppermost part of the DCM layer. The slight sub-optimal position of Synechococcus and prymnesiophytes with regard to their nutrient requirements suggests that their need for high irradiance plays a significant role in their location within the DCM. Finally, pelagophytes remained in deep layers without an apparent need for the high nutrient concentrations at those depths.

Epilimnetic production has declined in Lake Ontario, but increased production in metalimnetic deep chlorophyll layers (DCLs) may compensate for these losses. We investigated the spatial and temporal extent of DCLs, the mechanisms driving DCL formation, and the use of physical variables for predicting the depth and concentration of the deep chlorophyll maximum (DCM) during April–September 2013. A DCL with DCM concentrations 2 to 3 times greater than those in the epilimnion was present when the euphotic depth extended below the epilimnion, which occurred primarily from late June through mid-August. In situ growth was important for DCL formation in June and July, but settling and photoadaptation likely also contributed to the later-season DCL. Supporting evidence includes: phytoplankton biovolume was 2.4 × greater in the DCL than in the epilimnion during July, the DCL phytoplankton community of July was different from that of May and the July epilimnion ( p = 0.004), and there were concurrences of DCM with maxima in fine particle concentration and dissolved oxygen saturation. Higher nutrient levels in the metalimnion may also be a necessary condition for DCL formation because July metalimnetic concentrations were 1.5 × (nitrate) and 3.5 × (silica) greater than in the epilimnion. Thermal structure variables including epilimnion depth, thermocline depth, and thermocline steepness were useful for predicting DCM depth; the inclusion of euphotic depth only marginally improved these predictions. However, euphotic depth was critical for predicting DCM concentrations. The DCL is a productive and predictable feature of the Lake Ontario ecosystem during the stratified period. [ABSTRACT FROM AUTHOR]