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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]

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]

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]

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]

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]

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]

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]

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. [ABSTRACT FROM AUTHOR]

Photosynthetic O2 production can be an important source of oxygen in sub-surface ocean waters especially in permanently stratified oligotrophic regions of the ocean where O2 produced in deep chlorophyll maxima (DCM) is not likely to be outgassed. Today, permanently stratified regions extend across approximately 40% of the global ocean and their extent is expected to increase in a warmer ocean. Thus, predicting future ocean oxygen conditions requires a better understanding of the potential response of photosynthetic oxygen production to a warmer ocean. Based on our own and published observations of water column processes in oligotrophic regions, we develop a one-dimensional water column model describing photosynthetic oxygen production in the Sargasso Sea to quantify the importance of photosynthesis for the downward flux of O2 and examine how it may be influenced in a warmer ocean. Photosynthesis is driven in the model by vertical mixing of nutrients (including eddy-induced mixing) and diazotrophy and is found to substantially increase the downward O2 flux relative to physical-chemical processes alone. Warming (2°C) surface waters does not significantly change oxygen production at the DCM. Nor does a 15% increase in re-mineralization rate (assuming Q10 =2; 2°C warming) have significant effect on net sub-surface oxygen accumulation. However, changes in the relative production of particulate (POM) and dissolved organic material (DOM) generate relatively large changes in net sub-surface oxygen production. As POM/DOM production is a function of plankton community composition, this implies plankton biodiversity and food web structure may be important factors influencing O2 production in a warmer ocean. [ABSTRACT FROM AUTHOR]

At low trophic state, stable stratified water columns may provide favorable conditions for adapted phytoplankton species to form deep chlorophyll maxima (DCM). Such maxima occur regularly in Lake Stechlin, mainly contributed by the cyanobacterial species Cyanobium sp. and occasionally by Planktothrix rubescens. In the early twenty-first century, a rapid invasion by nostocalean cyanobacteria occurred in the lake and a number of Dolichospermum species together with Aphanizomenon flos- aquae appeared. As revealed by both microscopic and fluorimetric methods, during the summer stratification of 2013, a multispecific DCM was formed by Cyanobium, Planktothrix rubescens, and A. flos- aquae, however with spatial segregation. Planktothrix occurred in the upper hypolimnion, Aphanizomenon and Cyanobium dominated in the metalimnetic layer. Coexistence of these three cyanoprokaryota is possibly the consequence of different environmental factors limiting them (light, availability of N and P). This study represents a rare case when spatial niche segregation of phytoplankton species occurs in close to equilibrium conditions. DCM formed by Aphanizomenon and Cyanobium was detected by the fluoroprobe; Planktothrix with its different pigment compositions remained largely hidden. Our results indicate the necessity of parallel microscopic investigations and the need of careful calibration when fluorimetric methods are used for detecting cyanobacterial populations. [ABSTRACT FROM AUTHOR]

The community structure of planktonic cyanobacteria was studied in a dimictic lake in which recurrent summer surface algal blooms have frequently occurred since the beginning of this millennium. In eutrophic-hypereutrophic lakes, epilimnetic cyanobacterial blooms are promoted by increased ambient temperatures and water column thermal stability, which favour the vertical migration of buoyancyregulating cyanobacteria. Here we propose that intensified external energy (wind) that alters thermocline stability could explain the occurence of heavy blooms in the surface of lakes with low external nutrient loading. Specifically, we hypothesized that: i) in small stratified lakes with low external nutrient sources, cyanobacterial growth primarily occurs near the lake bottom, where phosphorus is more abundant and light is available; ii) we additionally hypothesized that turbulence induced by strong winds increases the amplitude and energy of metalimnetic internal waves and entrains meta- and hypolimnetic water, rich in nutrients and cyanobacteria, into the epilimnion. The study was done in a small lake (45 Ha, maximum and mean depth 7.2 m and 4.3 m, respectively) with mean epilimnetic dissolved phosphorus concentrations 4 µg L-1 and chlorophyll a 8 µg L-1. Vertical temperature profiles during the open season were continuously registered using thermistors. Weekly vertical profiles of light transmission, phytoplankton distribution and water chemistry were also taken. On one occasion, these variables were measured throughout a continuous 24 h cycle. Results demonstrated that summer cyanobacterial blooms were dominated by Plankthotrix spp., which began their cycle in late spring at the bottom of the lake, and grew to form dense metalimnetic biomass peaks. Time series analysis of isotherms and the Lake number indicated that internal metalimnetic waves (seiches) were present through the summer. During the diel sampling cycle, we found that medium to strong westerly wind gust events (~5 to >12 m s-1) induced large amplitude internal waves (mainly V2H1 mode) that vertically displaced the isotherms by more than 3.5 m. During this event the top of the metalimnetic algal peak was entrained through the epilimnion, bringing metalimnetic Plankthotrix spp. to the lake surface, modified the deep metalimnion and hypolimnion concentrations of dissolved oxygen, and caused an upsurge in phosphorus. We conclude that algal and nutrient upwelling linked to intermittent deep mixing events, play an important role in supporting summer cyanobacterial blooms in lake Bromont. [ABSTRACT FROM AUTHOR]

Iron limits carbon fixation in much of the modern ocean due to the very low solubility of ferric iron in oxygenated ocean waters. We examined iron-limitation of growth rate under varying light intensities in the coastal cyanobacterium Synechococcus bacillaris, a descendent of the oxygenic phototrophs that evolved ca. 3 billion years ago when the ocean was reducing and iron was present at much higher concentrations as soluble Fe(II). Decreasing light intensity increased the cellular iron:carbon (Fe:C) ratio needed to support a given growth rate, indicating that iron and light may co-limit the growth of Synechococcus in the ocean, as shown previously for eukaryotic phytoplankton. The cellular Fe:C ratios needed to support a given growth rate were 5- to 8-fold higher than ratios for coastal eukaryotic algae growing under the same light conditions. The higher iron requirements for growth in the coastal cyanobacterium may be largely caused by the high demand for iron in photosynthesis, and to higher ratios of iron-rich photosystem I to iron-poor photosystem II in Synechococcus than in eukaryotic algae. This high iron requirement may also be vestigial and represent an adaptation to the much higher iron levels in the ancient reducing ocean. Due to the high cellular iron requirement for photosynthesis and growth, and for low light acclimation, Synechococcus may be excluded from many low-iron and low-light environments. Indeed, it decreases rapidly with depth within the ocean's deep chlorophyll maximum (DCM) where iron and light levels are low, and lower-iron requiring picoeukaryotes typically dominate the biomass of phytoplankton community within the mid to lower DCM. [ABSTRACT FROM AUTHOR]

Deep chlorophyll maxima (DCM) are a common feature of poorly mixed, low-productivity lakes, but are less commonly observed in eutrophic systems. This study investigates the transition from a late-spring surface chlorophyll maximum (SCM) to an early summer DCM in a hypertrophic lake, identifying the phenology and mechanisms responsible for the transition and how the DCM was maintained in this hypertrophic system. Data and samples were collected weekly during summer and monthly in winter at five sampling stations in monomictic Lake Ōkaro, North Island, New Zealand. Additional samples were collected on two occasions at c. 3-5 h intervals over 24 h at a single station, corresponding to periods with a SCM and DCM, respectively. The mean DCM depth was correlated with mean thermocline depth, and euphotic depth ( z) continuously exceeded the depths of both the DCM and thermocline. The decline in the SCM and the transition to a DCM population were coincident with a gradual depletion of dissolved inorganic nutrients in surface waters and increased light penetration into the nutrient-rich hypolimnion. This study demonstrated that there are similarities between DCMs in lakes of high and low productivity, relating to the interplay between light, nutrients and thermal stratification. [ABSTRACT FROM AUTHOR]

The theory of resource competition in spatially extended systems with resources and biomass fluxes is far from trivial. Here, we analyze the competition between two phytoplankton species for light and a nutrient in a weakly mixed water column. We develop a general framework for such an analysis and show that the competition outcome can be largely understood from a single parameter, the slope of the invasion threshold in the plane of resources. Using this approach, we show that the competition outcome crucially depends on the depth of the biomass maximum. Under eutrophic conditions, when the phytoplankton production peaks on the surface, species composition depends on the ratio of resource supplies, and the competition outcome follows the "classic" rule: coexistence is possible if each competitor has the greatest effect on its most limiting resource. By contrast, in oligotrophic systems, characterized by deep biomass maxima, the absolute level of resource supplies drives species composition, and coexistence becomes more feasible if each competitor mostly consumes its least limiting resource. Finally, when the production peaks in the subsurface, good nutrient competitors are favored. Our findings are supported by empirical data. [ABSTRACT FROM AUTHOR]

The article discusses a study which investigated the sequencing of marine metagenomic fosmids that resulted in the discovery of new complete phage genomes. Topics discussed include the sequencing of a much larger set of fosmids, the existence of a biogeography of microbes, and the complexity of phage behavior. Also cited is the need to tap the largest genetic reservoir of the Earth using the advances in sequencing.

We deployed four Bio-Argo profiling floats in various oligotrophic locations of the Pacific subtropical gyres and Mediterranean Sea to address the seasonal phytoplankton dynamics in the euphotic layer and explore its dependence on light regime dynamics. Results show that there is a similar phytoplankton biomass seasonal pattern in the four observed oceanic regions. In the lower part of the euphotic layer, the seasonal displacement of the deep chlorophyll maximum (DCM) is light driven. During winter, the chlorophyll a concentration ([Chl a]) always increases in the upper euphotic mixed layer. This increase always results from a photoacclimation to the reduced irradiance. Depending on the location, however, the concentration can also be associated with an actual increase in biomass. The winter increase in [Chl a] results in an increase in irradiance attenuation that impacts the position of the isolume (level where the daily integrated photon flux is constant) and DCM, which becomes shallower. In summer when the [Chl a] in the upper layer decreases along with light attenuation, the DCM deepens and becomes closer to (and sometimes reaches) the nitracline, which enhances the phytoplankton biomass at the DCM. The bio-optical mechanisms and their relationship to light regimes that are revealed by the time series appear to be generic and potentially characteristic of all of the areas where a DCM forms, which is 50% of the open ocean. [ABSTRACT FROM AUTHOR]

Deep chlorophyll maxima (DCMs) are widespread features of oceans. In temperate regions, DCMs are commonly associated with isopycnal surfaces that frequently move over a wide vertical range. This general association between DCMs and isopycnals remains unexplained by present theories, and we show here that it emerges from the seasonal history of the water column. Analysis of the formation of more than 9000 seasonal DCMs throughout the world's oceans consistently locates the vertical position of spring/summer DCMs in temperate seas at the density of the previous winter mixed layer, independently of this density value and future depth. These results indicate that DCM formation cannot be understood without hysteresis by solely considering the instantaneous response of phytoplankton to vertical gradients in physical and chemical fields. Present theories for DCM formation cannot explain why spring and summer DCMs are systematically found at a density equal to that of the previous mixed layer where a bloom has occurred. Rather than reacting to instantaneous physical forcing, the results indicate that DCMs operate as self-preserving biological structures that are associated with particular isopycnals because of their capacity to modify the physicochemical environment. Combined with remote sensors to measure salinity and temperature in the surface ocean, this new understanding of DCM dynamics has the potential to improve the quantification of three-dimensional primary production via satellites. This significant enhancement of the representation of oceanic biological processes can also allow increasingly realistic predictions of future biogeochemical scenarios in a warming ocean. [ABSTRACT FROM AUTHOR]

Results from extensive cruises in the years 2000 and 2001 throughout distinct ecohydrodynamic regions of the central and southern North Sea are presented and used to generate estimates of gross primary production and new production. An undulating CTD fitted with a fluorometer was towed over a distance of 12,000 kms. Fluorescence data were used to determine the chlorophyll distribution and derive estimates of phytoplankton biomass. These results were combined with estimates of primary production (new and regenerated) from experiments from one cruise in order to estimate gross production for a greater geographical extent. Results from repeat inter-annual transects showed that the strength of the thermocline and the associated deep chlorophyll maximum were variable. However, when the primary production was integrated over the 15-40 m depth, the variability between years was low. While the depth and strength of the deep chlorophyll maximum varied across the region, a deep chlorophyll maximum (DCM) is a consistent and widespread feature of this region at around 30 m depth. In 2001 the calculated average primary production rate in summer for the whole area surveyed was 0.91 g C m day. This daily production equates to ~130 g C m for the summer stratified period. In the offshore stratified regions around the Dogger Bank and Eastern Central North Sea primary production of 64 g C m associated with the deep chlorophyll maximum (15-40 m) accounted for 60 % of total primary production during the summer stratified period (after the spring bloom). Approximately 66 % of new production in these areas occurred in the DCM. This study shows the extent of the DCM in the North Sea and demonstrates its importance in sustaining primary production after the spring bloom. [ABSTRACT FROM AUTHOR]

To better understand the vertical distribution of phytoplankton in the tropical and subtropical North Pacific, we used fast repetition rate fluorometry to investigate the photo-physiological condition of the phytoplankton assemblage in this region between February and March 2007. Along 155°E, between the equator and 24°N, the peak of fluorescence ( F), an indication of the deep chlorophyll maximum (DCM), was deeper than the top of the nitracline and occurred at the 2.4 ± 1.3 % (mean ± SD) light depth (relative to 0 m). The photochemical efficiency ( F/ F) and effective absorption cross-section of photosystem II (σ) were low at the surface but increased rapidly at depths between the top of the nitracline (40-138 m) and the DCM (70-158 m), an indication that the photo-physiological condition of the phytoplankton improved below the top of the nitracline. The depth of the maximal F/ F [Z( F/ F)] was 18-32 m deeper than the DCM and corresponded to the 0.8 ± 0.2 % light depth. The values of F/ F at the Z( F/ F) were 20 % higher than those at the DCM and averaged 0.48 ± 0.01. These results suggest that the phytoplankton assemblage beneath the DCM had a high potential photosynthetic performance capacity and was growing by using the very low ambient light in this region. [ABSTRACT FROM AUTHOR]

The cyanoprokaryote Planktothrix rubescens has been studied for over four decades in an alpine lake, Mondsee, Austria. We hypothesise that impacts of climate change can be distinguished from other environmental constraints controlling its population dynamics. During thermal stratification, P. rubescens formed a deep chlorophyll maximum in the metalimnion. Seasonal and diurnal depth distributions indicated that Planktothrix lived well below the euphotic zone, at low light levels and moderate temperatures. Photosynthetic parameters indicated a shade adapted population. The eutrophication and rehabilitation periods of Mondsee were characterised by fluctuations controlled by phosphorus, with periods below the P-threshold associated with low biovolumes and relatively stable populations. Positive net changes of Planktothrix biovolume occurred during the spring-summer transition and in autumn. During summer, the population usually declined to an annual minimum. The standardised residuals of annual biovolumes responded positively to the climate signal of the Winter North Atlantic Oscillation and the timing of the onset of stratification. An inverse relationship existed between off-set and persistence of stratification. P. rubescens only benefits from climate warming early in the year, during late spring overturn and early summer. Longer periods of summer stratification did not favour biovolume development. [ABSTRACT FROM AUTHOR]

In a medium-sized pre-alpine lake (North Italy) the cyanobacterium Planktothrix rubescens has strongly dominated the phytoplankton assemblage since 2000, similar to many pre-alpine lakes, despite improvements in water quality. The objective of this study was to determine the factors governing the spatial distribution of P. rubescens, including the major hydrodynamic processes and the influence of long-term reduction in nutrient concentrations during a period of climate warming. We used an intensive field campaign conducted from February 2010 to January 2011, to evaluate distributions of phytoplankton phyla, as well as P. rubescens, using spectrally resolved fluorescence measurements. These data provided highly spatially and temporally resolved phytoplankton population data suitable to calibrate and validate a coupled three-dimensional hydrodynamic (ELCOM) and ecological model (CAEDYM) of the lake ecosystem. The simulations revealed the fundamental role of physiological features of P. rubescens that led to observed vertical patterns of distribution, notably a deep chlorophyll maximum, and a strong influence of lake hydrodynamic processes, particularly during high-discharge inflows in summer stratification. The simulations are used to examine growth-limiting factors that help to explain the increased prevalence of P. rubescens during re-oligotrophication. [ABSTRACT FROM AUTHOR]

Resource competition in heterogeneous environments is still an unresolved problem of theoretical ecology. In this article, I analyze competition between two phytoplankton species in a deep water column, where the distributions of main resources (light and a limiting nutrient) have opposing gradients and co-limitation by both resources causes a deep biomass maximum. Assuming that the species have a trade-off in resource requirements and the water column is weakly mixed, I apply the invasion threshold analysis (Ryabov and Blasius, Ecol Lett 14:220-228, ) to determine relations between environmental conditions and phytoplankton composition. Although species deplete resources in the interior of the water column, the resource levels at the bottom and surface remain high. As a result, the slope of resources gradients becomes a new crucial factor which, rather than the local resource values, determines the outcome of competition. The value of resource gradients nonlinearly depend on the density of consumers. This leads to complex relationships between environmental parameters and species composition. In particular, it is shown that an increase of both the incident light intensity or bottom nutrient concentrations favors the best light competitors, while an increase of the turbulent mixing or background turbidity favors the best nutrient competitors. These results might be important for prediction of species composition in deep ocean. [ABSTRACT FROM AUTHOR]

profiling float equipped with a fluorimeter, a dissolved oxygen (DO) sensor, and temperature and salinity sensors was deployed in the subtropical mode water (STMW) formation region of the North Pacific. It acquired quasi-Lagrangian, 5-day-interval time-series records from March to July 2006. The time-series distribution of chlorophyll showed a sustained and sizable subsurface maximum at 50-100 m, just above the upper boundary of the STMW, throughout early summer (May-July). The DO concentration in this lower euphotic zone (50-100 m) was almost constant and supersaturated in the same period, becoming more supersaturated with time. On the other hand, the DO concentration at 100-150 m near the upper boundary of the STMW decreased much more slowly compared with the main layer of STMW below 150 m, even though oxygen consumption by organisms was expected to be larger in the former depth range. The small temporal variations of DO in the lower euphotic zone and near the upper boundary of the STMW were reasonably explained by downward oxygen transport because of large diapycnal diffusion near the top of the STMW. Assuming that the oxygen consumption rate at 100-150 m was the same as that in the main layer of STMW and compensated by the downward oxygen flux, the diapycnal diffusivity was estimated to be 1.7 × 10 m s. Nitrate transport into the euphotic zone by the same large diffusion was estimated to be 0.8 mmol N m day. All of the transported nitrate could have been used for photosynthesis by the phytoplankton; net community production was estimated to be 5.3 mmol C m day. [ABSTRACT FROM AUTHOR]

The vertical and seasonal distributions of the phytoflagellate Cryptomonas spp., and its most common, the planktonic ciliate predators ( Oligotrichida, Scuticociliatida, Hypotrichida and Prostomatida) were investigated in chemocline region of small saline, meromictic lake Shunet (Siberia, Russia) during 2003 and 2005. The lake has a pronounced chemocline, with abundance of purple and green sulphur bacteria. Vertical distribution of the Cryptomonas populations near the oxic/anoxic boundary layer was studied at close intervals in water sampled using a hydraulically operated thin-layer sampler. In both summer and winter, Cryptomonas peaked in water stratum 5–10 cm above anoxic zone or in the anoxic zone water column in the chemocline (about 5 m). Ciliate densities and biomass were also much higher in chemocline than in mixolimnion. The range of diurnal migration of Cryptomonas population was not very wide, and it was restricted to layers with high light intensity. The ciliates were sometimes detected above the upper border of the anoxic zone but also several centimetres below this zone. [ABSTRACT FROM AUTHOR]

The deep chlorophyll maximum (DCM) is a zone of maximal photosynthetic activity, generally located toward the base of the photic zone in lakes and oceans. In the tropical waters, this is a permanent feature, but in the Mediterranean and other temperate waters, the DCM is a seasonal phenomenon. The metagenome from a single sample of a mature Mediterranean DCM community has been 454 pyrosequenced both directly and after cloning in fosmids. This study is the first to be carried out at this sequencing depth (ca. 600 Mb combining direct and fosmid sequencing) at any DCM. Our results indicate a microbial community massively dominated by the high-light-adapted Prochlorococcus marinus subsp. pastoris, Synechococcus sp., and the heterotroph Candidatus Pelagibacter. The sequences retrieved were remarkably similar to the existing genome of P. marinus subsp. pastoris with a nucleotide identity over 98%. Besides, we found a large number of cyanophages that could prey on this microbe, although sequence conservation was much lower. The high abundance of phage sequences in the cellular size fraction indicated a remarkably high proportion of cells suffering phage lytic attack. In addition, several fosmids clearly belonging to Group II Euryarchaeota were retrieved and recruited many fragments from the total direct DNA sequencing suggesting that this group might be quite abundant in this habitat. The comparison between the direct and fosmids sequencing revealed a bias in the fosmid libraries against low-GC DNA and specifically against the two most dominant members of the community, Candidatus Pelagibacter and P. marinus subsp. pastoris, thus unexpectedly providing a feasible method to obtain large genomic fragments from other less prevalent members of this community. [ABSTRACT FROM AUTHOR]

In order to adequately assess the ecological status of thermally stratified lakes based on chlorophyll, the sampling must cover all productive layers of the water column. Missing the deep chlorophyll maxima (DCM) that often occur in the meta- or hypolimnion of transparent lakes supported by sufficient illumination and good nutrient availability may cause serious underestimation of the productivity and lead to misclassification of the lake ecological status. There is no commonly accepted sampling design for stratified lakes, and various monitoring guides suggest controversial designs. Our aim was to find some robust criteria to assess the probability of occurrence of a DCM and estimate the differences in measured mean chlorophyll concentrations caused by various sampling designs. Our theoretical model showed that the probability of occurrence of a DCM increases with increasing water transparency and decreasing lake size. Empirical data from Italian and Estonian stratified lakes confirmed the results. Testing of different sampling designs on lakes with full measured chlorophyll profiles available showed that taking only surface layer samples will lead with a high probability to an underestimation of the chlorophyll concentration in the trophogenic layer. In order not to miss the Chl peak in stratified lakes, in most cases it would be more precautious not to limit the sampling to the well-mixed epilimnion but to extend it to the whole euphotic layer. Sampling the epilimnion instead of the euphotic zone could cause up to a 70% underestimation of the chlorophyll concentration, an error that would cause a misclassification of the lake by one or even two status classes in a 5-class assessment system. In most cases, the 2.5 * Secchi depths proved a suitable criterion of the sampling depth and only in the case of surface scums, would sampling of a 3 * Secchi depth layer be recommended in order not to miss the deep chlorophyll maximum. [ABSTRACT FROM AUTHOR]

The factors affecting vertical distributions of chlorophyll fluorescence were examined in four temperate, warm monomictic lakes. Each of the lakes (maximum depth >80 m) was sampled over 2 years at intervals from monthly to seasonal. Profiles were taken of chlorophyll fluorescence (as a proxy for algal biomass), temperature and irradiance, as well as integrated samples from the surface mixed layer for chlorophyll a (chl a) and nutrient concentrations in each lake. Depth profiles of chlorophyll fluorescence were also made along transects of the longest axis of each lake. Chlorophyll fluorescence maxima occurred at depths closely correlated with euphotic depth ( r2 = 0.67, P < 0.01), which varied with nutrient status of the lakes. While seasonal thermal density stratification is a prerequisite for the existence of a deep chlorophyll maximum (DCM), our study provides evidence that the depth of light penetration largely dictates the DCM depth during stratification. Reduction in water clarity through eutrophication can cause a shift in phytoplankton distributions from a DCM in spring or summer to a surface chlorophyll maximum within the surface mixed layer when the depth of the euphotic zone ( zeu) is consistently shallower than the depth of the surface mixed layer ( zSML). Trophic status has a key role in determining vertical distributions of chlorophyll in the four lakes, but does not appear to disrupt the annual cycle of maximum chlorophyll in winter. [ABSTRACT FROM AUTHOR]

Phytoplankton compete for nutrients and light in a vertically heterogeneous environment determined by turbulent mixing. We analyzed a model of competition between two phytoplankton species in a stratified water column. We assume that the surface layer is uniformly mixed and that the deep layer is poorly mixed, as is commonly observed in lakes and oceans. We employed two analytical techniques, Iout- R theory in the mixed surface layer and a game theoretical approach in the deep layer. Under our assumptions, at equilibrium, each species is either absent or resides in the benthic layer, the deep layer, or the surface layer. Assuming a trade-off between nutrient- and light-competitive abilities, we obtained five spatial configurations of coexistence and the corresponding parameter regions where they occur. Good light competitors show two distinct ecological niches: in mesotrophic conditions, they live as understory species below a layer of good nutrient competitors; in eutrophic conditions, they live as competitive dominants in the surface layer. Multiple regions of alternative stable states are possible in parameter space. This work extends previous phytoplankton competition theory to stratified water columns, as commonly found in lakes and oceans. [ABSTRACT FROM AUTHOR]

Deep chlorophyll a maximum (DCM) was found in the 80-110mdeep northeastern coastal Adriatic (Pag and Velebit Channels) in July 2007. Eight times higher concentrations of chlorophyll were determined in the 50 to 60 m DCM layer (0.81 to 0.91 mg L-1) relative to the surface waters. According to thermohaline conditions and phytoplankton biomass distribution, over the greater part of the considered area the water column was separated into three layers (the surface layer approximately 10 m deep, the subsurface layer extending between 10 and 50m, and the deep layer found below 50mdepth) and three distinctive regimes were detected (in the South Velebit Channel, the Middle Velebit Channel, and the Pag Channel). This is the first evidence of the deep chlorophyll maximum in the eastern coastal Adriatic Sea. Deep chlorophyll maxima were composed of picoplanktonic, rounded cells, probably cyanobacteria (1.2 mm in size, 8 × 103 - 1 × 104 cells mL-1). [ABSTRACT FROM AUTHOR]

A coupled 1D physical-biological model of Crater Lake is presented. The model simulates the seasonal evolution of two functional phytoplankton groups, total chlorophyll, and zooplankton in good quantitative agreement with observations from a 10-year monitoring study. During the stratified period in summer and early fall the model displays a marked vertical structure: the phytoplankton biomass of the functional group 1, which represents diatoms and dinoflagellates, has its highest concentration in the upper 40 m; the phytoplankton biomass of group 2, which represents chlorophyta, chrysophyta, cryptomonads and cyanobacteria, has its highest concentrations between 50 and 80 m, and phytoplankton chlorophyll has its maximum at 120 m depth. A similar vertical structure is a reoccurring feature in the available data. In the model the key process allowing a vertical separation between biomass and chlorophyll is photoacclimation. Vertical light attenuation (i.e., water clarity) and the physiological ability of phytoplankton to increase their cellular chlorophyll-to-biomass ratio are ultimately determining the location of the chlorophyll maximum. The location of the particle maxima on the other hand is determined by the balance between growth and losses and occurs where growth and losses equal. The vertical particle flux simulated by our model agrees well with flux measurements from a sediment trap. This motivated us to revisit a previously published study by Dymond et al. (). Dymond et al. used a box model to estimate the vertical particle flux and found a discrepancy by a factor 2.5–10 between their model-derived flux and measured fluxes from a sediment trap. Their box model neglected the exchange flux of dissolved and suspended organic matter, which, as our model and available data suggests is significant for the vertical exchange of nitrogen. Adjustment of Dymond et al.’s assumptions to account for dissolved and suspended nitrogen yields a flux estimate that is consistent with sediment trap measurements and our model. [ABSTRACT FROM AUTHOR]

Based on the in situ optical measurements in the Bohai Sea of China, which belongs to a typical case-2 water area, we studied the characteristics of DCM (deep chlorophyll maximum) such as its spatial distribution, vertical profile, etc. We found that when the depth of the chlorophyll maximum is comparatively small, even in turbid coastal water regions, there is always a good correlation between the concentrations of chlorophyll maximum and the satellite-received signals in blue-green spectral bands; the correlation is even better than that between the surface chlorophyll concentrations and the satellite-received signals. The strong correlation existing even in turbid coastal water regions indicates that an ocean color model to retrieve the concentration of DCM can be constructed for coastal waters if a comprehensive knowledge of the vertical distribution of chlorophyll concentration in the Bohai Sea of China is available. [ABSTRACT FROM AUTHOR]

Vertical distribution of phytoplankton and the formation of deep chlorophyll maximum (DCM) in the metalimnion of a small stratified and partly meromictic temperate lake was studied in 1999 and 2000. During summer DCM usually occurred on the borderline of H2S and oxygen-containing waters. At the depths where the bacteriochlorophyll (Bchl) maxima were observed, the sulphide concentration was usually relatively low compared to the bottom layers, where its concentration reached as high as possible saturation level. In April 2000, DCM was formed at the depth of 3.5 m, and lowered thereafter slowly to 6.5 m by October. The concentration of Bchl d reached the highest values (over 1000 μg l−1) just before the water column was mixed up in autumn. In December and April Bchl d was detectable only near the bottom of the lake. The concentration of chlorophyll a yielded by the spectrophotometric phaeopigment corrected method and by HPLC (high pressure liquid chromatography), fit rather well in the upper layers. In deeper water layers chlorophyll a concentration (Chl a) measured by spectrophotometry was overestimated about 47 times if compared to HPLC values because of the high Bchl d in that layer. In most cases vertical profiles of primary production (PP) did not coincide with the vertical distribution of the pigment content; the maximum values of PP were found in the epilimnion. In some cases PP had notably high values also at the depth of DCM. In the upper layers Chl a usually did not exceeded 20 μg l−1 in spring and 10 μg l−1 in summer. The moderately high Chl a in the epilimnion in spring was significantly reduced after the formation of thermocline most probably because of the establishment of the nutrient limitation in epilimnion. Decreasing Chl a concentration in the epilimnion led to increased water transparency and better light conditions for photosynthetic bacteria in metalimnion. [ABSTRACT FROM AUTHOR]

1. The unicellular green alga Chlamydomonas acidophila accumulates in a thin phytoplankton layer in the hypolimnion (deep chlorophyll maximum, DCM) of an extremely acidic lake (Lake 111, pH 2.6, Lusatia, Germany), in which the underwater light spectrum is distorted and red-shifted. 2. Chlamydomonas acidophila exhibited a significantly higher absorption efficiency and a higher cellular chlorophyll b content when incubated in the red shifted underwater light of Lake 111 than in a typical, blue-green dominated, light spectrum. 3. Chlamydomonas acidophila has excellent low light acclimation properties (increased chlorophyll b content, increased oxygen yield and a low light saturation point for photosynthesis) that support survival of the species in the low light climate of the DCM. 4. In situ acclimation to the DCM under low light and temperature decreased maximum photosynthetic rate in autotrophic C. acidophila cultures, whereas the presence of glucose under these conditions enhanced photosynthetic efficiency and capacity. 5. The adaptive abilities of C. acidophila to light and temperature shown in this study, in combination with the absence of potent competitors because of low lake pH, most probably enable the unusual dominance of the green alga in the DCM of Lake 111. [ABSTRACT FROM AUTHOR]

Allozyme diversity of a population of D. pulex was studied over two seasons in a small lake (Gräfenhain, Saxony, Germany). In this eutrophic lake, accumulation of hydrogen sulfide during winter ice covers leads to an annual killing-off of cladocerans. Hence, the population must be re-established every year by hatchlings from sexual eggs. The absence of planktivorous fish due to strong biomanipulation ensures high abundances of Daphnia from June until September. Besides the investigation of seasonal changes of genotype diversity, vertical differences between individuals from the epilimnion and from a deep chlorophyll maximum (DCM) were studied. Specimens were classified by allozyme electrophoresis using six allozyme loci. Except for aldehyde oxidase (AO) all tested allozyme loci were monomorphic. Two alleles were found at the AO-locus in the overall study, allowing the population to be subdivided into three clonegroups (ss, fs, ff). The proportion of each clonegroup both in the epilimnion and in the DCM did not vary over the two seasons and no differences were found in the year-to-year comparison. Although two completely different habitats (epilimnion, DCM) were found to be used by D. pulex, clonal structure in both depth strata was similar at all times indicating no intraspecific segregation of the population. In spite of a non-permanent population with periodical sexual reproduction, a constant deviation from Hardy-Weinberg-equilibrium was detected in the overall study. This remarkable genetic structure is interpreted to be the result of both founder effects and a strong and long-lasting selection pressure caused by 15 years of stationary conditions due to intense biomanipulation. [ABSTRACT FROM AUTHOR]

Water mass distributions, upwelling and filament formation are of complex significance for biological and chemical processes. Temperature and salinity are routinely used to characterize these hydrographic conditions. In situ profiles and sample analyses from the cruise So119 of RV Sonne in May 1997 into the Arabian Sea indicate that optical parameters such as the fluorescence of phytoplankton pigments, gelbstoff and proteins reflect hydrographic processes as well, and are useful to describe and explain biochemical processes. The dependence of bio-optical parameters to DOC and chlorophyll a concentrations is examined. While the relation between extracted chlorophyll and chlorophyll fluorescence is quite reasonable, DOC and gelbstoff fluorescence do not show a functional dependency. This is mainly due to sunlight-induced degradation of fluorophores in the surface layer. Enhanced gelbstoff signals in intermediate and deep waters are related to Persian Gulf Water. The optical data reveal a relation between the depth of the chlorophyll maximum, the depth of 1% surface light and the geographical latitude. In the context of remote sensing of phytoplankton biomass from ocean colour images, the depth of the chlorophyll maximum is a crucial parameter. Thus, the revealed dependency on latitude is useful for the evaluation of primary production with satellite remote sensing. [ABSTRACT FROM AUTHOR]

SUMMARY 1. In oligotrophic lakes, phytoplankton and bacteria growing in the deep chlorophyll maximum in the cool metalimnion of lakes often dominate biomass and production, but the importance of this source of food for zooplankton is unknown. 2. During much of the day, Daphnia rosea in two mountain lakes inhabited deep chlorophyll layers where food availability was at least equal to that in the epilimnion. 3. To determine the importance of the two strata (epilimnion and metalimnion) for Daphnia , we used a cross-classified factorial experiment to measure how epilimnetic and metalimnetic food and temperature (10 and 16 °C) influenced survival, growth and reproduction. 4. Daphnia survived and grew better when fed seston from the epilimnion of one lake, although chlorophyll, particulate nitrogen and particulate carbon were 2–2.5 times greater in the metalimnion. 5. Temperature had no significant influence on Daphnia survival or growth. Similar results were obtained with food from the second lake, with Daphnia surviving and reproducing better when provided with epilimnetic, rather than metalimnetic food, although the quantities of chlorophyll and carbon in the two strata were similar. 6. Food quality, rather than quantity or temperature, appeared to be the most important determinant influencing survival, growth and reproduction, and the greater food quantity in the metalimnia was not used effectively by the Daphnia . [ABSTRACT FROM AUTHOR]

The microstratification of the microbial community at the chemocline of Lake Cadagno and the associated inorganic carbon fixation activity was studied by fine layer sampling. A deep chlorophyll maximum caused by diatoms overlying Cryptomonas was found at the upper edge of the chemocline. A high population density of phototrophic sulphur bacteria, mainly Amoebobacter cf. purpureus, occurred closely below the oxic-anoxic boundary. Despite the small fraction of total lake volume represented by the chemocline, half of the total carbon photoassimilation of the lake occurred within the chemocline with approximately equal contributions by oxygenic and anoxygenic phototrophs. Rates of dark carbon fixation in the chemocline were even higher than rates of photoassimilation, especially at the depths where oxygen and sulphide coexisted during part of the day. These results indicate a substantial contribution by chemolithotrophic organisms to the carbon cycle in Lake Cadagno. Analysis of stable carbon isotopes suggests that zooplankton may obtain as much as half of its carbon at the chemocline, indicating a strong link between production in anoxic waters and the food web in the oxic part of the lake. [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 CO 2 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 R 2 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 FROM AUTHOR]

Clones of Cryptomonas phaseolus Skuja, Cryptonuonas rostratiformis (Skuja) Skuja in Hubber-Pestalozzi, and Cryptomonas undulata Gervais were isolated from the deep chlorophyll maximum near the oxu/anoxic boundary layer of the mesoeutrophic lake Schlachtensee, Germany Different autecological features of these species were studied in batch culture experiments. Cryptomonas cf. ovata Ehvenberg and Chroomonas sp. That never dominated in the deep chlorophyll layer were also isolated from Schlachtensee to study their light-dependent growth in comparison to the deep-living species. Cryptomonas undulata. C. cf. ovata, and C. Phaseolus had a very low light compensation point (5-7 μ mol-m[sup-2]-s[sup-1]), whereas the growth rate of Chroomonas sp. and C. rostratiformis was positive above 16 and 24 μ mol-m[sup-2]-s[sup-1], Cryptomonas phaseolus and Chroomonas sp. Became photoinhibited above photon flux densities of 92 and 116 μ mol-m[sup-2]-s[sup-1], Cryptomonas rostratiformis, C. cf. ovata, and C. undulata reached a maximum growth rate at a considerably higher photon flux density (198-250 μ mol-m[sup-2]-s[sup-1]). Cryptomonas phaseolus grew fastest under light-limiting conditions. Cryptpmonas phaseolus and C. undulata were best able to survive prolonged periods of darkness, Cryptomonas phaseolus, C. Rostratiformis, and C. undulata did not show any uptake of fluorescent latex beads. When labeled glucose was provided in naturally occurring concentrations, carbon uptake by C. phaseolus, C. rostratiformis, and C undulata was negligibly small in comparison to cellular carbon content. I suggest that the adaption to a low-light environment is an important preadaption for the dominance of C. phaseolus and C. undulata near the freshwater chemocline. [ABSTRACT FROM AUTHOR]