ALBERT

Description: We examined the total diatom flux and species composition, total coccolith flux and total mass flux collected with a sediment trap between October 1993 and January 2006 in the northeastern entrance of the Gulf of Lions (North Western Mediterranean). The average daily diatom and coccolith fluxes (3 x 10 7 valves m 2 d –1 and 6.1 x 10 8 coccoliths m 2 day –1 , respectively) are comparable in magnitude with previously reported data sets in other high productivity areas of the Western Mediterranean. The temporal particle flux pattern reflected the variations in surface oceanographic conditions and primary productivity, which showed strong annual cycling. Highest diatom, coccolith and total mass fluxes always occurred during the winter–spring transition, while minima were observed during summer. Changes in the diverse diatom communities reflected the water column conditions throughout the record. The intensity of the diatom winter–spring blooms seemed to be enhanced in those years with intense and cold winds during winter, whereas years with low winter wind stress were liable to be less productive for diatoms. Coccolith fluxes exhibited a more stable interannual pattern than diatom fluxes. Significant discrepancies were found between the sediment trap and surficial sediment diatom assemblages.

Description: The coccolithophore Emiliania huxleyi is a marine phytoplankton species capable of forming small calcium carbonate scales (coccoliths) which cover the organic part of the cell. Calcification rates of E. huxleyi are known to be sensitive to changes in seawater carbonate chemistry. It is, however, not yet understood how these changes are reflected in the morphology of coccoliths. Here, we compare data on coccolith size, weight, and malformation from a~set of five experiments with a large diversity of carbonate chemistry conditions. This diversity allows distinguishing the influence of individual carbonate chemistry parameters such as carbon dioxide (CO2), bicarbonate (HCO3−), carbonate (CO32−), and protons (H+) on the measured parameters. Measurements of fine-scale morphological structures reveal an increase of coccolith malformation with decreasing pH suggesting that H+ is the major factor causing malformations. Coccolith distal shield area varies from about 5 to 11 μm2. Changes in size seem to be mainly induced by varying [HCO3−] and [H+] although influence of [CO32−] cannot be entirely ruled out. Changes in coccolith weight were proportional to changes in size. Increasing CaCO3 production rates are reflected in an increase in coccolith weight and an increase of the number of coccoliths formed per unit time. The combined investigation of morphological features and coccolith production rates presented in this study may help to interpret data derived from sediment cores, where coccolith morphology is used to reconstruct calcification rates in the water column.

Description: Ocean acidification is a result of the uptake of anthropogenic CO2 from the atmosphere into the ocean and has been identified as a major environmental and economic threat. The release of several thousands of petagrams of carbon over a few hundred years will overwhelm the capacity of the surface ocean reservoirs to absorb carbon. The recorded and anticipated changes in seawater carbonate chemistry will presumably affect the global oceanic carbonate production. Coccolithophores as the primary calcifying phytoplankton group, and especially Emiliania huxleyi as the most abundant species have shown a reduction of calcification at increased CO2 concentrations for the majority of strains tested in culture experiments. A reduction of calcification is associated with a decrease in coccolith weight. However, the effect in monoclonal cultures is relatively small compared to the strong variability displayed in natural E. huxleyi communities, as these are a mix of genetically and sometimes morphologically distinct types. Average coccolith weight is likely influenced by the variability in seawater carbonate chemistry in different parts of the worlds' oceans and on glacial/interglacial time scales due to both physiological effects and morphotype selectivity. An effect of the ongoing ocean acidification on E. huxleyi calcification has so far not been documented in situ. Here, we analyze E. huxleyi coccolith weight from the NW Mediterranean Sea in a 12 yr sediment trap series, and surface sediment and sediment core samples using an automated recognition and analyzing software. Our findings clearly show (1) a continuous decrease in the average coccolith weight of E. huxleyi from 1993 to 2005, reaching levels below pre-industrial Holocene and industrial 20th century values recorded in the sedimentary record, and (2) seasonal variability in coccolith weight that is linked to the coccolithophore production. The observed long-term decrease in coccolith weight is most likely a result of the changes in the surface ocean carbonate system. Our results provide first indications of an in situ impact of ocean acidification on coccolithophore weight in a natural E. huxleyi population even in the highly alkaline Mediterranean Sea.

Description: Coccolith calcification is known to respond to ocean acidification in culture experiments as well as in present and past oceans. Previous studies basically focus on changes in coccolith weight due to increasing CO2 and the resulting changes in the carbonate system but pay little attention to the influence of other environmental factors. In order to untangle changes in coccolithophore calcification due to environmental factors such as temperature and/or productivity from changes caused by increasing pCO2 and carbonate ion concentration we here present a study on coccolith calcification from the Holocene North Atlantic Ocean. The pre-industrial Holocene with its predominantly stable carbonate system provides the conditions for such a comprehensive analysis. For a realistic analysis on changes in major components of Holocene coccolithophores, the family Noelaerhabdaceae was selected, which constitutes the main part of the assemblage in the North Atlantic. Records of average coccolith weights from three Holocene sediment cores along a North–South transect in the North Atlantic were analysed. During the Holocene mean weight (and therefore calcification) of Noelaerhabdaceae (E. huxleyi and Gephyrocapsa) coccoliths decreases at the Azores (Geofar KF 16) from around 7 to 5.5 pg, but increases at the Rockall Plateau (ODP Site 980) from around 6 to 8 pg and at the Vøring Plateau (MD08-3192) from 7 to 10.5 pg. This amplitude of average weight variability is within the range of glacial/interglacial changes that were interpreted to be an effect of decreasing carbonate ion concentration. By comparison with SEM assemblage counts, we show that weight changes are partly due to variations in the coccolithophore assemblage, but also an effect of a change in calcification and/or morphotype variability within single species. Our results indicate that there is no single key factor responsible for the observed changes in coccolith weight. A major increase in coccolith weight occurs during a slight decrease in carbonate ion concentration in the Late Holocene at the Rockall Plateau and Vøring Plateau. Here, more favourable productivity conditions apparently lead to an increase in coccolith weight, either due to the capability of coccolithophore species, especially E. huxleyi, to adapt to decreasing carbonate ion concentration, or due to a shift towards heavier calcifying morphotypes.

Description: The coccolithophore Emiliania huxleyi is a marine phytoplankton species capable of forming small calcium carbonate scales (coccoliths) which cover the organic part of the cell. Calcification rates of E. huxleyi are known to be sensitive to changes in seawater carbonate chemistry. It has, however, not yet been clearly determined how these changes are reflected in size and weight of individual coccoliths and which specific parameter(s) of the carbonate system drive morphological modifications. Here, we compare data on coccolith size, weight, and malformation from a set of five experiments with a large diversity of carbonate chemistry conditions. This diversity allows distinguishing the influence of individual carbonate chemistry parameters such as carbon dioxide (CO2), bicarbonate (HCO3−), carbonate ion (CO32−), and protons (H+) on the measured parameters. Measurements of fine-scale morphological structures reveal an increase of coccolith malformation with decreasing pH suggesting that H+ is the major factor causing malformations. Coccolith distal shield area varies from about 5 to 11 μm2. Changes in size seem to be mainly induced by varying [HCO3−] and [H+] although influence of [CO32−] cannot be entirely ruled out. Changes in coccolith weight were proportional to changes in size. Increasing CaCO3 production rates are reflected in an increase in coccolith weight and an increase of the number of coccoliths formed per unit time. The combined investigation of morphological features and coccolith production rates presented in this study may help to interpret data derived from sediment cores, where coccolith morphology is used to reconstruct calcification rates in the water column.

Description: The relevance of biological Si cycling for dissolved silica (DSi) export from terrestrial biogeosystems is still in debate. Even in systems showing a high content of weatherable minerals, like Cambisols on volcanic tuff, biogenic Si (BSi) might contribute 〉 50% to DSi (Gerard et al., 2008). However, the number of biogeosystem studies is rather limited for generalized conclusions. To cover one end of controlling factors on DSi, i.e., weatherable minerals content, we studied a forested site with absolute quartz dominance (〉 95%). Here we hypothesise minimal effects of chemical weathering of silicates on DSi. During a four year observation period (05/2007–04/2011), we quantified (i) internal and external Si fluxes of a temperate-humid biogeosystem (beech, 120 yr) by BIOME-BGC (version ZALF), (ii) related Si budgets, and (iii) Si pools in soil and beech, chemically as well as by SEM-EDX. For the first time two compartments of biogenic Si in soils were analysed, i.e., phytogenic and zoogenic Si pool (testate amoebae). We quantified an average Si plant uptake of 35 kg Si ha−1 yr−1 – most of which is recycled to the soil by litterfall – and calculated an annual biosilicification from idiosomic testate amoebae of 17 kg Si ha−1. The comparatively high DSi concentrations (6 mg L−1) and DSi exports (12 kg Si ha−1 yr−1) could not be explained by chemical weathering of feldspars or quartz dissolution. Instead, dissolution of a relictic, phytogenic Si pool seems to be the main process for the DSi observed. We identified canopy closure accompanied by a disappearance of grasses as well as the selective extraction of pine trees 30 yr ago as the most probable control for the phenomena observed. From our results we concluded the biogeosystem to be in a transient state in terms of Si cycling.

Description: Ocean acidification is a result of the uptake of anthropogenic CO2 from the atmosphere into the ocean and has been identified as a major environmental and economic threat. The release of several thousands of petagrams of carbon over a few hundred years will have an overwhelming effect on surface ocean carbon reservoirs. The recorded and anticipated changes in seawater carbonate chemistry will presumably affect global oceanic carbonate production. Coccolithophores as the primary calcifying phytoplankton group, and especially Emiliania huxleyi as the most abundant species have shown a reduction of calcification at increased CO2 concentrations for the majority of strains tested in culture experiments. A reduction of calcification is associated with a decrease in coccolith weight. However, the effect in monoclonal cultures is relatively small compared to the strong variability displayed in natural E. huxleyi communities, as these are a mix of genetically and sometimes morphologically distinct types. Average coccolith weight is likely influenced by the variability in seawater carbonate chemistry in different parts of the world's oceans and on glacial/interglacial time scales due to both physiological effects and morphotype selectivity. An effect of the ongoing ocean acidification on E. huxleyi calcification has so far not been documented in situ. Here, we analyze E. huxleyi coccolith weight from the NW Mediterranean Sea in a 12-year sediment trap series, and surface sediment and sediment core samples using an automated recognition and analyzing software. Our findings clearly show (1) a continuous decrease in the average coccolith weight of E. huxleyi from 1993 to 2005, reaching levels below pre-industrial (Holocene) and industrial (20th century) values recorded in the sedimentary record and (2) seasonal variability in coccolith weight that is linked to the coccolithophore productivity. The observed long-term decrease in coccolith weight is most likely a result of the changes in the surface ocean carbonate system. Our results provide the first indications of an in situ impact of ocean acidification on coccolithophore weight in a natural E. huxleyi population, even in the highly alkaline Mediterranean Sea.

Description: The response of coccolithophore calcification to ocean acidification has been studied in culture experiments as well as in present and past oceans. The response, however, is different between species and strains, and for the relatively small carbonate chemistry changes observed in natural environments, a uniform response of the entire coccolithophore community has not been documented so far. Moreover, previous palaeo-studies basically focus on changes in coccolith weight due to increasing CO2 and the resulting changes in the carbonate system, and only few studies focus on the influence of other environmental factors. In order to untangle changes in coccolithophore calcification due to environmental factors such as temperature and/or productivity from changes caused by increasing pCO2 and decreasing carbonate ion concentration, we here present a study on coccolith calcification from the Holocene North Atlantic Ocean. The pre-industrial Holocene, with its predominantly stable atmospheric CO2, provides the conditions for such a comprehensive analysis. For an analysis on changes in major components of Holocene coccolithophores under natural conditions, the family Noelaerhabdaceae was selected, which constitutes the main part of the assemblage in the North Atlantic. Records of average coccolith weights from three Holocene sediment cores along a north–south transect in the North Atlantic were analysed. During the Holocene, mean weight (and therefore calcification) of Noelaerhabdaceae (Emiliania huxleyi and Gephyrocapsa) coccoliths decreased at the Azores (Geofar KF 16) from around 7 to 6 pg, but increased at the Rockall Plateau (ODP site 980) from around 6 to 8 pg, and at the Vøring Plateau (MD08-3192) from 7 to 10 pg. The amplitude of average weight variability is within the range of glacial–interglacial changes that were interpreted to be an effect of decreasing carbonate ion concentration. By comparison with SEM assemblage counts, we show that weight changes are not only partly due to variations in the coccolithophore assemblage but also an effect of a change in calcification and/or morphotype variability within single species. Our results indicate that there is no single key factor responsible for the observed changes in coccolith weight. A major increase in coccolith weight occurs during a slight decrease in carbonate ion concentration in the late Holocene at the Rockall Plateau and Vøring Plateau. Here, more favourable productivity conditions apparently lead to an increase in coccolith weight, either due to the capability of coccolithophore species, especially E. huxleyi, to adapt to decreasing carbonate ion concentration or due to a shift towards heavier calcifying morphotypes.