Alkenone and foraminifera records from sediment samples off Namibia

Radiocarbon stratigraphy is an essential tool for high resolution paleoceanographic studies. Age models based on radiocarbon ages of foraminifera are commonly applied to a wide range of geochemical studies, including the investigation of temporal leads and lags. The critical assumption is that temporal coupling between foraminifera and other sediment constituents, including specific molecular organic compounds (biomarkers) of marine phytoplankton, e.g. alkenones, is maintained in the sediments. To test this critical assumption in the Benguela upwelling area, we have determined radiocarbon ages of total C37-C39 alkenones in 20 samples from two gravity cores and three multicorer cores. The cores were retrieved from the continental shelf and slope off Namibia, and samples were taken from Holocene, deglacial and Last Glacial Maximum core sections. The alkenone radiocarbon ages were compared to those of planktic foraminifera, total organic carbon, fatty acids and fine grained carbonates from the same samples. Interestingly, the ages of alkenones were 1000 to 4500 yr older than those of foraminifera in all samples. Such age differences may be the result of different processes: Bioturbation associated with grain size effects, lateral advection of (recycled) material and redeposition of sediment on upper continental slopes due to currents or tidal movement are examples for such processes. Based on the results of this study, the age offsets between foraminifera and alkenones in sediments from the upper continental slope off Namibia most probably do not result from particle-selective bioturbation processes. Resuspension of organic particles in response to tidal movement of bottom waters with velocities up to 25 cm/s recorded near the core sites is the more likely explanation. Our results imply that age control established using radiocarbon measurements of foraminifera may be inadequate for the interpretation of alkenone-based proxy data. Observed temporal leads and lags between foraminifera based data and data derived from alkenone measurements may therefore be secondary signals, i.e. the result of processes associated with particle settling and biological activity.

(Appendix) Carbon and nitrogen isotope ratios of ODP Site 160-964

Nitrogen isotopes of chlorins, degradation products of chlorophyll, reflect the isotopic composition of nutrient N utilized by marine phytoplankton communities. Here we show that in sediments of the eastern Mediterranean Pleistocene and Holocene, values of d15N for chlorins and total nitrogen vary in concert, with a consistent offset of ~5 per mil reflecting the fractionation imparted during chlorophyll biosynthesis. Samples from the Integrated Ocean Drilling Program Sites 964 and 969 were analyzed at a sampling resolution of ~4-10 cm, clustered around sapropel events 2, 3, 4 and 5 (~100-170 ka). In low organic content sediments, chlorin values of ~0 per mil coincident with total nitrogen values of ~+ 5 per mil indicate that the latter reflects the original biomass and is not a consequence of diagenetic isotope enrichment. In sapropel horizons, the chlorin and total nitrogen values are 5 per mil more negative (~-5 per mil and ~ 0 per mil, respectively), resembling previously-reported, modern-day water-column particulates (~0 per mil). We suggest that nutrient conditions in the Eastern Mediterranean correspond to three scenarios and that the similarity between sapropel and modern-day bulk d15N is coincidental. Organic-poor marl sediments formed under oligotrophic conditions where surface productivity resulted from upwelling of Atlantic-sourced nitrate. Sapropels were characterized by enhanced diazotrophy that was likely fueled by increased riverine P fluxes to surface waters. Present-day conditions are dominated by anthropogenic N sources. These scenarios agree with a model of sapropel formation in which stratification caused by increased fresh-water inputs led to N fixation due to P:N nutrient imbalance. Enhanced production combined with stratification promoted and maintained anoxic deep waters, consequently increasing organic matter preservation. Such a model may be relevant to interpreting other episodes of intense organic matter deposition in past oceans.

Experiment: Response of Nodularia spumigena to pCO2

The filamentous and diazotrophic cyanobacterium Nodularia spumigena plays a major role in the productivity of the Baltic Sea as it forms extensive blooms regularly. Under phosphorus limiting conditions Nodularia spumigena has a high enzyme affinity for dissolved organic phosphorus (DOP) by production and release of alkaline phosphatase. Additionally, it is able to degrade proteinaceous compounds by expressing the extracellular enzyme leucine aminopeptidase. As atmospheric CO2 concentrations are increasing, we expect marine phytoplankton to experience changes in several environmental parameters including pH, temperature, and nutrient availability. The aim of this study was to investigate the combined effect of CO2-induced changes in seawater carbonate chemistry and of phosphate deficiency on the exudation of organic matter, and its subsequent recycling by extracellular enzymes in a Nodularia spumigena culture. Batch cultures of Nodularia spumigena were grown for 15 days aerated with three different pCO2 levels corresponding to values from glacial periods to future values projected for the year 2100. Extracellular enzyme activities as well as changes in organic and inorganic compound concentrations were monitored. CO2 treatment-related effects were identified for cyanobacterial growth, which in turn was influencing exudation and recycling of organic matter by extracellular enzymes. Biomass production was increased by 56.5% and 90.7% in the medium and high pCO2 treatment, respectively, compared to the low pCO2 treatment and simultaneously increasing exudation. During the growth phase significantly more mucinous substances accumulated in the high pCO2 treatment reaching 363 µg Gum Xanthan eq /l compared to 269 µg Gum Xanthan eq /l in the low pCO2 treatment. However, cell-specific rates did not change. After phosphate depletion, the acquisition of P from DOP by alkaline phosphatase was significantly enhanced. Alkaline phosphatase activities were increased by factor 1.64 and 2.25, respectively, in the medium and high compared to the low pCO2 treatment. In conclusion, our results suggest that Nodularia spumigena can grow faster under elevated pCO2 by enhancing the recycling of organic matter to acquire nutrients.

A role for diatom-like silicon transporters in calcifying coccolithophores.

Biomineralization by marine phytoplankton, such as the silicifying diatoms and calcifying coccolithophores, plays an important role in carbon and nutrient cycling in the oceans. Silicification and calcification are distinct cellular processes with no known common mechanisms. It is thought that coccolithophores are able to outcompete diatoms in Si-depleted waters, which can contribute to the formation of coccolithophore blooms. Here we show that an expanded family of diatom-like silicon transporters (SITs) are present in both silicifying and calcifying haptophyte phytoplankton, including some globally important coccolithophores. Si is required for calcification in these coccolithophores, indicating that Si uptake contributes to the very different forms of biomineralization in diatoms and coccolithophores. Significantly, SITs and the requirement for Si are absent from highly abundant bloom-forming coccolithophores, such as Emiliania huxleyi. These very different requirements for Si in coccolithophores are likely to have major influence on their competitive interactions with diatoms and other siliceous phytoplankton.

A role for diatom-like silicon transporters in calcifying coccolithophores.

Biomineralization by marine phytoplankton, such as the silicifying diatoms and calcifying coccolithophores, plays an important role in carbon and nutrient cycling in the oceans. Silicification and calcification are distinct cellular processes with no known common mechanisms. It is thought that coccolithophores are able to outcompete diatoms in Si-depleted waters, which can contribute to the formation of coccolithophore blooms. Here we show that an expanded family of diatom-like silicon transporters (SITs) are present in both silicifying and calcifying haptophyte phytoplankton, including some globally important coccolithophores. Si is required for calcification in these coccolithophores, indicating that Si uptake contributes to the very different forms of biomineralization in diatoms and coccolithophores. Significantly, SITs and the requirement for Si are absent from highly abundant bloom-forming coccolithophores, such as Emiliania huxleyi. These very different requirements for Si in coccolithophores are likely to have major influence on their competitive interactions with diatoms and other siliceous phytoplankton.

Studies on phytoplankton of the estuarine and marine environments

This thesis deals with the results oi’ investigations on phytoplankton productivity and related aspects conducted in various ecoaystms such as estuarine, inshore and oceanic enviroments and certain special ecosytans including the pu.-awn culture fields and associated many-eves, mud bank and the seas around the Andaman-Nicobar Islands. This study also includes the qualitative and quantitative variations of phytoplankton production, their seasonal abundance, factors controlling the same and the magnitude of the potential resources derived 1!:-om it

Effects of solar radiation on marine bacterial and phytoplankton heterotrophic activities

Programa de doctorado: Oceanografía (Bienio 2006-2008). Universidad de Las Palmas de Gran Canaria, Departamento de Biología y Institut de Ciéncies del Mar.

Increased carbon dioxide availability alters phytoplankton stoichiometry and affects carbon cycling and growth of a marine planktonic herbivore

Rising levels of CO2 in the atmosphere have led to increased CO2 concentrations in the oceans. This enhanced carbon availability to the marine primary producers has the potential to change their nutrient stoichiometry, and higher carbon to nutrient ratios are expected. As a result, the quality of the primary producers as food for herbivores may change. Here, we present experimental work showing the effect of feeding Rhodomonas salina grown under different pCO2 (200, 400 and 800 µatm) on the copepod Acartia tonsa. The rate of development of copepodites decreased with increasing CO2 availability to the algae. The surplus carbon in the algae was excreted by the copepods, with younger stages (copepodites) excreting most of their surplus carbon through respiration, and adult copepods excreting surplus carbon mostly as DOC. We consider the possible consequences of different excretory pathways for the ecosystem. A continued increase in the CO2 availability for primary production, together with changes in the nutrient loading of coastal ecosystems, may cause changes in the trophic links between primary producers and herbivores.

Limiting factors for phytoplankton growth in subtropical reservoirs: the effect of light and nutrient availability in different longitudinal compartments

Limited information is available on the interactions between environmental factors and algal growth in tropical and subtropical aquatic systems. We investigated the relationships between algal biomass (measured as chlorophyll, Chl-a) and light, total phosphorus (TP) and total nitrogen (TN) in longitudinal zones of subtropical reservoirs. We studied the seasonal variation of water variables in Itupararanga Reservoir (Brazil) and compared the results with 16 other subtropical lakes and reservoirs. The longitudinal zones in Itupararanga Reservoir were considered statistically different (p 0.05, MANOVA). From the riverine zone to the dam region of the reservoir, Spearman Correlation Test suggested that light limitation and TP limitation tended to decrease and increase, respectively. Although nitrate concentrations were high (400 g/L), the Spearman coefficients between Chl-a and TN and the TN:TP ratios (11:1 TN:TP 35:1) indicated that nitrogen may be co-limiting algal growth in the studied water body. Putting Itupararanga in a regional context allowed assessment of potential influences of land use on trophic state. Within the subtropical dataset, TP explained a greater percentage of variance in Chl-a (R2 = 0.70) than TN (R2 = 0.17). The main land use type within the reservoirs drainage area significantly influenced the concentrations of TP, TN, and Chl-a (p 0.05, MANOVA), with different relationships between nutrients and chlorophyll in forested (R2 = 0.12-0.33), agricultural (R2 = 0.50-0.68) and urban (R2 = 0.09-0.64) watersheds. Comparisons with literature values and those from reservoirs with less altered watersheds indicated that Itupararanga Reservoir is reaching the mesotrophic-eutrophic boundary, and further nutrient enrichment could cause water quality degradation.

Effects of concentrated viral communities on photosynthesis and community composition of co-occurring benthic microalgae and phytoplankton

Marine viruses have been shown to affect phytoplankton productivity; however, there are no reports on the effect of viruses on benthic microalgae (microphytobenthos). Hence, this study investigated the effects of elevated concentrations of virus-like particles on the photosynthetic physiology and community composition of benthic microalgae and phytoplankton. Virus populations were collected near the sediment surface and concentrated by tangential flow ultrafiltration, and the concentrate was added to benthic and water column samples that were obtained along a eutrophication gradient in the Brisbane River/Moreton Bay estuary, Australia. Photosynthetic and community responses of benthic microalgae, phytoplankton and bacteria were monitored over 7 d in aquaria and in situ. Benthic microalgal communities responded to viral enrichment in both eutrophic and oligotrophic sediments. In eutrophic sediments, Euglenophytes (Euglena sp.) and bacteria decreased in abundance by 20 to 60 and 26 to 66%, respectively, from seawater controls. In oligotrophic sediments, bacteria decreased in abundance by 30 to 42% from seawater controls but the dinoflagellate Gymnodinium sp. increased in abundance by 270 to 3600% from seawater controls, The increased abundance of Gymnodinium sp. may be related to increased availability of dissolved organic matter released from lysed bacteria. Increased (140 to 190% from seawater controls) initial chlorophyll a fluorescence measured with a pulse-amplitude modulated fluorometer was observed in eutrophic benthic microalgal incubations following virus enrichment, consistent with photosystem II damage. Virus enrichment in oligotrophic water significantly stimulated carbon fixation rates, perhaps due to increased nutrient availability by bacterial lysis. The interpretation of data from virus amendment experiments is difficult due to potential interaction with unidentified bioactive compounds within seawater concentrates. However, these results show that viruses are capable of influencing microbial dynamics in sediments.