Marine Primary Productivity Is Driven by a Selection Effect

10 páginas, 6 figuras, 1 tabla.-- Pedro Cermeño ... et al.-- This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these termsThe number of species of autotrophic communities can increase ecosystem productivity through species complementarity or through a selection effect which occurs when the biomass of the community approaches the monoculture biomass of the most productive species. Here we explore the effect of resource supply on marine primary productivity under the premise that the high local species richness of phytoplankton communities increases resource use through transient selection of productive species. Using concurrent measurements of phytoplankton community structure, nitrate fluxes into the euphotic zone, and productivity from a temperate coastal ecosystem, we find that observed productivities are best described by a population growth model in which the dominant species of the community approach their maximum growth rates. We interpret these results as evidence of species selection in communities containing a vast taxonomic repertory. The prevalence of selection effect was supported by open ocean data that show an increase in species dominance across a gradient of nutrient availability. These results highlight the way marine phytoplankton optimize resources and sustain world food stocks. We suggest that the maintenance of phytoplankton species richness is essential to sustain marine primary productivity since it guarantees the occurrence of highly productive speciesThis research was supported by grants CTM2011-25035, CTM2012-30680, and CGL2013-41256-P from the Spanish Government (SG). PC and SV are supported by Ramon y Cajal contracts from the SGPeer reviewe

La ecología del plancton en la oceanografía biológica: un homenaje a la labor de Marta Estrada

Plankton ecology has been the object of intense research and progress in the last few decades. This has been partly due to technological advances that have facilitated the multidisciplinary and high-resolution sampling of ecosystems and improved experimentation and analytical methodologies, and to sophisticated modelling. In addition, exceptional researchers have had the vision to integrate all these innovative tools to form a solid theoretical background in ecology. Here we provide an overview of the outstanding research work conducted by Professor Marta Estrada and her pioneering contribution to different areas of research in the last four decades. Her research in biological oceanography has mainly focussed on phytoplankton ecology, taxonomy and physiology, the functional structure of plankton communities, and physical and biological interactions in marine ecosystems. She has combined a variety of field and laboratory approaches and methodologies, from microscopy to satellite observations, including in-depth statistical data analysis and modelling. She has been a reference for scientists all over the world. Here, her contributions to plankton ecology are summarized by some of her students and closest collaborators, who had the privilege to share their science and everyday experiences with her.La ecología del plancton ha experimentado una intensa investigación y progreso en las últimas décadas. Esta se ha debido, en parte, a los avances tecnológicos que han facilitado la toma de muestras multidisciplinar y de alta resolución de los ecosistemas, la mejora de la experimentación y metodologías analíticas y el modelado matemático/numérico. El papel de investigadores excepcionales que han tenido la visión de integrar todas estas herramientas innovadoras a una sólida formación teórica en la ecología ha sido clave en este avance. A continuación hemos elaborado un resumen de la tarea de investigación llevada a cabo por la profesora Marta Estrada y su contribución pionera a diferentes áreas de investigación en las últimas cuatro décadas. Su investigación, en el marco de la oceanografía biológica, se ha centrado principalmente en la ecología del fitoplancton, su taxonomía y fisiología, la estructura funcional de las comunidades de plancton y las interacciones físicas y biológicas en los ecosistemas marinos. Ha combinado una variedad de enfoques y metodologías de campo y de laboratorio, desde observaciones por satélite, hasta el análisis en profundidad de datos estadísticos y el uso del modelado tanto analítico como numérico. La profesora Estrada ha sido una referencia para los científicos alrededor del mundo. Aquí, su contribución a la ecología del plancton la resumen algunos de sus alumnos y colaboradores más cercanos que han tenido el privilegio de compartir ciencia y vivencias con ella

Uncoupled seasonal variability of transparent exopolymer and Coomassie stainable particles in coastal Mediterranean waters: Insights into sources and driving mechanisms

Transparent exopolymer particles (TEP) and Coomassie stainable particles (CSP) are gel-like particles, ubiquitous in the ocean, that affect important biogeochemical processes including organic carbon cycling by planktonic food webs. Despite much research on both groups of particles (especially TEP) over many years, whether they exist as distinctly stainable fractions of the same particles or as independent particles, each with different driving factors, remains unclear. To address this question, we examined the temporal dynamics of TEP and CSP over 2 complete seasonal cycles at 2 coastal sites in the Northwestern Mediterranean Sea, the Blanes Bay Microbial Observatory (BBMO) and the L’Estartit Oceanographic Station (EOS), as well as their spatial distribution along a coast-to-offshore transect. Biological, chemical, and physical variables were measured in parallel. Surface concentrations (mean + standard deviation [SD]) of TEP were 36.7 + 21.5 µg Xanthan Gum (XG) eq L–1 at BBMO and 36.6 + 28.3 µg XG eq L–1 at EOS; for CSP, they were 11.9 + 6.1 µg BSA eq L–1 at BBMO and 13.0 + 5.9 µg BSA eq L–1 at EOS. Seasonal variability was more evident at EOS, where surface TEP and CSP concentrations peaked in summer and spring, respectively, and less predictable at the shore-most station, BBMO. Vertical distributions between surface and 80 m, monitored at EOS, showed highest TEP concentrations within the surface mixed layer during the stratification period, whereas CSP concentrations were highest before the onset of summer stratification. Phytoplankton were the main drivers of TEP and CSP distributions, although nutrient limitation and saturating irradiance also appeared to play important roles. The dynamics and distribution of TEP and CSP were uncoupled both in the coastal sites and along the transect, suggesting that they are different types of particles produced and consumed differently in response to environmental variability

Los aportes de aerosoles afectan las propiedades ópticas de la materia orgánica disuelta en las aguas costeras del Mediterráneo Noroccidental

10 pages, 6 figures, 1 table[EN] Aeolian inputs of organic and inorganic nutrients to the ocean are important as they can enhance biological production in surface waters, especially in oligotrophic areas like the Mediterranean. The Mediterranean littoral is particularly exposed to both anthropogenic and Saharan aerosol depositions on a more or less regular basis. During the last few decades experimental studies have been devoted to examining the effect of inorganic nutrient inputs from dust on microbial activity. In this study, we performed experiments at two different locations of the NW Mediterranean, where we evaluated the changes in the quality and quantity of dissolved organic matter due to atmospheric inputs of different origin (Saharan and anthropogenic) and its subsequent transformations mediated by microbial activities. In both experiments the humic-like and protein-like substances, and the fluorescence quantum yield increased after addition. In general, these changes in the quality of dissolved organic matter did not significantly affect the prokaryotes. The recalcitrant character of the fluorescent dissolved organic matter (FDOM) associated with aerosols was confirmed, as we found negligible utilization of chromophoric compounds over the experimental period. We framed these experiments within a two-year time series data set of atmospheric deposition and coastal surface water analyses. These observations showed that both Saharan and anthropogenic inputs induced changes in the quality of organic matter, increasing the proportion of FDOM substances. This increase was larger during Saharan dust events than in the absence of Saharan influence[ES] Los aportes atmosféricos de nutrientes orgánicos e inorgánicos al océano son importantes ya que pueden aumentar la producción biológica en aguas superficiales, especialmente en las zonas oligotróficas como el Mediterráneo. El litoral del Mediterráneo está particularmente expuesto a aportes de origen antropogénico y a deposiciones de polvo sahariano de forma más o menos regular. Durante las últimas décadas los estudios experimentales se han dedicado, sobre todo, a examinar el efecto de la entrada de nutrientes inorgánicos atmosféricos sobre la actividad microbiana. En este estudio, se realizaron experimentos con comunidades microbianas procedentes de dos zonas del Mediterráneo noroccidental. Se evaluaron los cambios en la calidad y cantidad de la materia orgánica disuelta debido a aportes atmosféricos de distinto origen y sus posteriores transformaciones mediadas por actividades microbianas. En ambos experimentos las sustancias orgánicas fluorescentes y el rendimiento cuántico de fluorescencia aumentaron después de la adición de material atmosférico. En general, estos cambios en la calidad de la materia orgánica no afectaron significativamente a los organismos procariotas. El carácter recalcitrante de la materia orgánica disuelta fluorescente (FDOM) contenida en los aerosoles se confirmó ya que la utilización de compuestos cromóforos durante el período experimental fue insignificante. Los resultados obtenidos se contextualizan en relación con una serie temporal de dos años de datos adquiridos de deposición atmosférica y análisis de agua superficial costera. La variabilidad temporal de estas dos variables mostró que tanto los aportes saharianos como antropogénicos provocaron cambios en la calidad de la materia orgánica disuelta en aguas superficiales, incrementando la fracción fluorescente. Éste aumento resultó ser mayor durante eventos de polvo sahariano que en ausencia de ellosThis study was supported by the projects 1179-1198. ADEPT (CTM2011-23458), DOREMI (CTM2012-342949) and ANIMA (CTM2015-65720-R MINECO/FEDER, UE). E.D. Sánchez-Pérez would like to thank the Consejo Nacional de Ciencia y Tecnologia (CONACyT) for their financial support through a PhD fellowship. I. Marín thanks the FPI Spanish scholarship programme for its support (BES 2012-052976)Peer Reviewe

Diatom Dominance Enhances Resistance of Phytoplanktonic POM to Mesopelagic Microbial Decomposition

Particulate organic matter (POM) lability is one of the key factors determining the residence time of organic carbon (OC) in the marine system. Phytoplankton community composition can influence the rate at which heterotrophic microorganisms decompose phytoplankton detrital particles and thus, it controls the fraction of OC that reaches the ocean depths, where it can be sequestered for climate-relevant spans of time. Here, we compared the degradation dynamics of POM from phytoplankton assemblages of contrasting diatom dominance in the presence of mesopelagic prokaryotic communities during a 19-day degradation experiment. We found that diatom-derived POM exhibited an exponential decay rate approximately three times lower than that derived from a community dominated by flagellated phytoplankton (mainly coccolithophores and nanoflagellates). Additionally, dissolved organic matter (DOM) released during the degradation of diatom particles accumulated over the experiment, whereas only residual increases in DOM were detected during the degradation of non-diatom materials. These results suggest that diatom-dominance enhances the efficiencies of the biological carbon pump and microbial carbon pump through the relatively reduced labilities of diatom particles and of the dissolved materials that arise from their microbial processing

Viral-mediated microbe mortality modulated by ocean acidification and eutrophication: Consequences for the carbon fluxes through the microbial food web

Anthropogenic carbon emissions are causing changes in seawater carbonate chemistry including a decline in the pH of the oceans. While its aftermath for calcifying microbes has been widely studied, the effect of ocean acidification (OA) on marine viruses and their microbial hosts is controversial, and even more in combination with another anthropogenic stressor, i.e., human-induced nutrient loads. In this study, two mesocosm acidification experiments with Mediterranean waters from different seasons revealed distinct effects of OA on viruses and viral-mediated prokaryotic mortality depending on the trophic state and the successional stage of the plankton community. In the winter bloom situation, low fluorescence viruses, the most abundant virus-like particle (VLP) subpopulation comprising mostly bacteriophages, were negatively affected by lowered pH with nutrient addition, while the bacterial host abundance was stimulated. High fluorescence viruses, containing cyanophages, were stimulated by OA regardless of the nutrient conditions, while cyanobacteria of the genus Synechococcus were negatively affected by OA. Moreover, the abundance of very high fluorescence viruses infecting small haptophytes tended to be lower under acidification while their putative hosts\u27 abundance was enhanced, suggesting a direct and negative effect of OA on viral–host interactions. In the oligotrophic summer situation, we found a stimulating effect of OA on total viral abundance and the viral populations, suggesting a cascading effect of the elevated pCO2 stimulating autotrophic and heterotrophic production. In winter, viral lysis accounted for 30 ± 16% of the loss of bacterial standing stock per day (VMMBSS) under increased pCO2 compared to 53 ± 35% in the control treatments, without effects of nutrient additions while in summer, OA had no significant effects on VMMBSS (35 ± 20% and 38 ± 5% per day in the OA and control treatments, respectively). We found that phage production and resulting organic carbon release rates significantly reduced under OA in the nutrient replete winter situation, but it was also observed that high nutrient loads lowered the negative effect of OA on viral lysis, suggesting an antagonistic interplay between these two major global ocean stressors in the Anthropocene. In summer, however, viral-mediated carbon release rates were lower and not affected by lowered pH. Eutrophication consistently stimulated viral production regardless of the season or initial conditions. Given the relevant role of viruses for marine carbon cycling and the biological carbon pump, these two anthropogenic stressors may modulate carbon fluxes through their effect on viruses at the base of the pelagic food web in a future global change scenario

Los aportes de aerosoles afectan las propiedades ópticas de la materia orgánica disuelta en las aguas costeras del Mediterráneo Noroccidental

Aeolian inputs of organic and inorganic nutrients to the ocean are important as they can enhance biological production in surface waters, especially in oligotrophic areas like the Mediterranean. The Mediterranean littoral is particularly exposed to both anthropogenic and Saharan aerosol depositions on a more or less regular basis. During the last few decades experimental studies have been devoted to examining the effect of inorganic nutrient inputs from dust on microbial activity. In this study, we performed experiments at two different locations of the NW Mediterranean, where we evaluated the changes in the quality and quantity of dissolved organic matter due to atmospheric inputs of different origin (Saharan and anthropogenic) and its subsequent transformations mediated by microbial activities. In both experiments the humic-like and protein-like substances, and the fluorescence quantum yield increased after addition. In general, these changes in the quality of dissolved organic matter did not significantly affect the prokaryotes. The recalcitrant character of the fluorescent dissolved organic matter (FDOM) associated with aerosols was confirmed, as we found negligible utilization of chromophoric compounds over the experimental period. We framed these experiments within a two-year time series data set of atmospheric deposition and coastal surface water analyses. These observations showed that both Saharan and anthropogenic inputs induced changes in the quality of organic matter, increasing the proportion of FDOM substances. This increase was larger during Saharan dust events than in the absence of Saharan influence.Los aportes atmosféricos de nutrientes orgánicos e inorgánicos al océano son importantes ya que pueden aumentar la producción biológica en aguas superficiales, especialmente en las zonas oligotróficas como el Mediterráneo. El litoral del Mediterráneo está particularmente expuesto a aportes de origen antropogénico y a deposiciones de polvo sahariano de forma más o menos regular. Durante las últimas décadas los estudios experimentales se han dedicado, sobre todo, a examinar el efecto de la entrada de nutrientes inorgánicos atmosféricos sobre la actividad microbiana. En este estudio, se realizaron experimentos con comunidades microbianas procedentes de dos zonas del Mediterráneo noroccidental. Se evaluaron los cambios en la calidad y cantidad de la materia orgánica disuelta debido a aportes atmosféricos de distinto origen y sus posteriores transformaciones mediadas por actividades microbianas. En ambos experimentos las sustancias orgánicas fluorescentes y el rendimiento cuántico de fluorescencia aumentaron después de la adición de material atmosférico. En general, estos cambios en la calidad de la materia orgánica no afectaron significativamente a los organismos procariotas. El carácter recalcitrante de la materia orgánica disuelta fluorescente (FDOM) contenida en los aerosoles se confirmó ya que la utilización de compuestos cromóforos durante el período experimental fue insignificante. Los resultados obtenidos se contextualizan en relación con una serie temporal de dos años de datos adquiridos de deposición atmosférica y análisis de agua superficial costera. La variabilidad temporal de estas dos variables mostró que tanto los aportes saharianos como antropogénicos provocaron cambios en la calidad de la materia orgánica disuelta en aguas superficiales, incrementando la fracción fluorescente. Éste aumento resultó ser mayor durante eventos de polvo sahariano que en ausencia de ellos

Horizontal and Vertical Distributions of Transparent Exopolymer Particles (TEP) in the NW Mediterranean Sea Are Linked to Chlorophyll a and O2 Variability

12 pages, 6 figures, 3 tables, supplementary material http://journal.frontiersin.org/article/10.3389/fmicb.2016.02159/full#supplementary-materialTransparent Exopolymer Particles (TEP) are relevant in particle and carbon fluxes in the ocean, and have economic impact in the desalination industry affecting reverse osmosis membrane fouling. However, general models of their occurrence and dynamics are not yet possible because of the poorly known co-variations with other physical and biological variables. Here, we describe TEP distributions in the NW Mediterranean Sea during late spring 2012, along perpendicular and parallel transects to the Catalan coast. The stations in the parallel transect were sampled at the surface, while the stations in the perpendicular transect were sampled from the surface to the bathypelagic, including the bottom nepheloid layers. We also followed the short-term TEP dynamics along a 2-day cycle in offshore waters. TEP concentrations in the area ranged from 4.9 to 122.8 and averaged 31.4 ± 12.0 μg XG eq L−1. The distribution of TEP measured in transects parallel to the Catalan Coast correlated those of chlorophyll a (Chla) in May but not in June, when higher TEP-values with respect to Chla were observed. TEP horizontal variability in epipelagic waters from the coast to the open sea also correlated to that of Chla, O2 (that we interpret as a proxy of primary production) and bacterial production (BP). In contrast, the TEP vertical distributions in epipelagic waters were uncoupled from those of Chla, as TEP maxima were located above the deep chlorophyll maxima. The vertical distribution of TEP in the epipelagic zone was correlated with O2 and BP, suggesting combined phytoplankton (through primary production) and bacterial (through carbon reprocessing) TEP sources. However, no clear temporal patterns arose during the 2-day cycle. In meso- and bathypelagic waters, where phytoplanktonic sources are minor, TEP concentrations (10.1 ± 4.3 μg XG eq l−1) were half those in the epipelagic, but we observed relative TEP increments coinciding with the presence of nepheloid layers. These TEP increases were not paralleled by increases in particulate organic carbon, indicating that TEP are likely to act as aggregating agents of the mostly inorganic particles present in these bottom nepheloid layersThis work was funded by projects funded by the Spanish Ministry of Science STORM (CTM2009-09352/MAR), SUMMER (CTM2008-03309/MAR), DOREMI (CTM2012-34294), REMEI (CTM2015-70340-R), ANIMA (CTM2015-65720-R), PEGASO (CTM2012-37615), and Grup consolidat de Recerca de la Generalitat de Catalunya (2014SGR/1179)Peer Reviewe

Regularidades estacionales en la producción primaria y los parámetros fotosintéticos en una estación costera del NO Mediterráneo

We carried out monthly photosynthesis-irradiance (P-E) experiments with the 14C-method for 12 years (2003–2014) to determine the photosynthetic parameters and primary production of surface phytoplankton in the Blanes Bay Microbial Observatory, a coastal sampling station in the NW Mediterranean Sea. Our goal was to obtain seasonal trends and to establish the basis for detecting future changes of primary production in this oligotrophic area. The maximal photosynthetic rate PBmax ranged 30-fold (0.5-15 mg C mg Chl a–1 h–1), averaged 3.7 mg C mg Chl a–1 h–1 (±0.25 SE) and was highest in August and lowest in April and December. We only observed photoinhibition twice. The initial or light-limited slope of the P-E relationship, αB, was low, averaging 0.007 mg C mg Chl a–1 h–1 (μmol photons m–2 s–1)–1 (±0.001 SE, range 0.001-0.045) and showed the lowest values in spring (April-June). The light saturation parameter or saturation irradiance, EK, averaged 711 μmol photons m–2 s–1 (± 58.4 SE) and tended to be higher in spring and lower in winter. Phytoplankton assemblages were typically dominated by picoeukaryotes in early winter, diatoms in late autumn and late winter, dinoflagellates in spring and cyanobacteria in summer. Total particulate primary production averaged 1.45 mg C m-3 h–1 (±0.13 SE) with highest values in winter (up to 8.50 mg C m-3 h–1) and lowest values in summer (summer average, 0.30 mg C m-3 h–1), while chlorophyll-specific primary production averaged 2.49 mg C mg Chl a–1 h–1 (±0.19, SE) and peaked in summer (up to 12.0 mg C mg Chl a–1 h–1 in August). 14C-determined phytoplankton growth rates varied between ca. 0.3 d–1 in winter and 0.5 d–1 in summer and were within 60-80% of the maximal rates of growth, based on PBmax. Chlorophyll a was a good predictor of primary production only in the winter and autumn. Seasonality appeared to explain most of the variability in the studied variables, while phytoplankton composition played a minor role. Daily integrated primary production was fairly constant throughout the year: similar to previous oxygen-based estimates in winter but considerably lower than these in summer. The difference between 14C- and oxygen-based estimates of primary production could be explained by community respiration. Annually integrated primary production amounted to a rather modest 48 g C m–2 yr–1 (equivalent to 130 mg C m–2 d–1). Although no interannual patterns were detected, our work soundly establishes the seasonal trends for the coastal NW Mediterranean, therefore setting the basis for future detection of change.Hemos llevado a cabo mediciones mensuales de la relación entre fotosíntesis e irradiancia (curvas P-E) con el método del 14C durante 12 años (2003-2014), para obtener los parámetros fotosintéticos y la producción primaria del fitoplancton superficial en el Observatorio Microbiano de la Bahía de Blanes, una estación de muestreo costera en el noroeste del Mediterráneo. Nuestro objetivo era determinar las tendencias estacionales y establecer la línea de base para detectar cambios futuros de la producción primaria en esta área oligotrófica. La tasa fotosintética máxima PBmaxosciló 30 veces (0.5 a 15 mg C mg Chl a–1 h–1), con un promedio de 3.7 mg C mg Chl a–1 h–1 (±0.25, error estándar), y fue máxima en agosto y mínima en abril y diciembre. Sólo se observó fotoinhibicion dos veces. La pendiente inicial de la curva P-E, αB, fue baja, con un promedio de 0.007 mg C mg Chl a–1 h–1 (µmol fotones m–2 s–1)–1 (error estándar ±0.001, rango de 0.001-0.045) y presentó los valores más bajos en primavera (abril-junio). El parámetro de saturación de irradiancia, EK, presentó un promedio de 711 µmol fotones m–2 s–1 (±58.4, error estándar) y tendió a ser mayor en primavera y menor en invierno. El fitoplancton estuvo típicamente dominado por picoeucariotas a principios de invierno, diatomeas a finales de otoño y en el invierno avanzado, dinoflagelados en primavera, y cianobacterias en verano. La producción primaria particulada total promedió 1.45 m–3 h–1 (±0.13 error estándar) con máximos en invierno (hasta 8.50 mg C m–3 h–1) y mínimos en verano (media en verano, 0.30 mg C m–3 h–1), mientras que la producción primaria por unidad de clorofila promedió 2.49 mg C mg Chl a–1 h–1 (±0.19, error estándar) y alcanzó su punto máximo en verano (hasta 12.0 mg C mg Chl a–1 h–1 en agosto). Las tasas de crecimiento del fitoplancton determinadas por 14C oscilaron entre ca. 0.3 d–1 en invierno y 0.5 d–1 en verano, siendo 60-80% de las tasas máximas de crecimiento basadas en los valores de PBmax. La clorofila a resultó ser un buen predictor de la producción primaria sólo en el invierno y el otoño. La estacionalidad explica la mayor parte de la variabilidad en las variables estudiadas, mientras que la composición del fitoplancton juega un papel menor. La producción primaria integrada diaria fue bastante constante durante todo el año, similar a estimaciones anteriores con mediciones de oxígeno en invierno, pero considerablemente más bajas en verano. La diferencia entre las estimaciones de la producción primaria por 14C y las estimaciones en base al oxígeno podrían explicarse por la respiración de la comunidad. Anualmente la producción primaria integrada ascendió a valores más bien modestos de 48 g C m–2 año–1 (equivalentes a 130 mg C m–2 d–1). Aunque no se detectaron patrones interanuales, nuestro trabajo establece las tendencias estacionales para la producción primaria en el litoral Mediterráneo noroccidental, y supone la línea de base para la detección de cambios futuros