Uma investigação das sequências de fase Hebbianas descritas como grafos de assembleias neuronais

Hebb proposed that synapses between neurons that fire synchronously are strengthened, forming cell assemblies and phase sequences. The former, on a shorter scale, are ensembles of synchronized cells that function transiently as a closed processing system; the latter, on a larger scale, correspond to the sequential activation of cell assemblies able to represent percepts and behaviors. Nowadays, the recording of large neuronal populations allows for the detection of multiple cell assemblies. Within Hebb’s theory, the next logical step is the analysis of phase sequences. Here we detected phase sequences as consecutive assembly activation patterns, and then analyzed their graph attributes in relation to behavior. We investigated action potentials recorded from the adult rat hippocampus and neocortex before, during and after novel object exploration (experimental periods). Within assembly graphs, each assembly corresponded to a node, and each edge corresponded to the temporal sequence of consecutive node activations. The sum of all assembly activations was proportional to firing rates, but the activity of individual assemblies was not. Assembly repertoire was stable across experimental periods, suggesting that novel experience does not create new assemblies in the adult rat. Assembly graph attributes, on the other hand, varied significantly across behavioral states and experimental periods, and were separable enough to correctly classify experimental periods (Naïve Bayes classifier; maximum AUROCs ranging from 0.55 to 0.99) and behavioral states (waking, slow wave sleep, and rapid eye movement sleep; maximum AUROCs ranging from 0.64 to 0.98). Our findings agree with Hebb’s view that neuronal assemblies correspond to primitive building blocks of representation, nearly unchanged in 10 the adult, while phase sequences are labile across behavioral states and change after novel experience. The results are compatible with a role for phase sequences in behavior and cognition

(Table 2) Zinc complexation data from POLARSTERN cruise ANT-XXIV/3

The speciation of dissolved zinc (Zn) was investigated by voltammetry in the Atlantic sector of the Southern Ocean along two transects across the major frontal systems: along the Zero Meridian and across the Drake Passage. In the Southern Ocean south of the APF we found detectable labile inorganic Zn throughout the surface waters in contrast to studies from lower latitudes. Using a combination of ASV titrations and pseudopolarography revealed the presence of significant concentration of electrochemically inert Zn ligands throughout the Southern Ocean. These ligands however were nearly always saturated due to the presence of excess concentrations of dissolved Zn that were associated with the high nutrient waters south of the Antarctic Polar Front (APF). Only in surface waters did the concentration of Zn complexing ligands exceed the dissolved Zn concentrations suggesting a biological source for these ligands. Our findings have clear implications for the biogeochemical cycling of Zn and for the interpretation of paleo records utilizing Zn in opal as a tracer of Zn speciation in the water column.

(Table 2) Ba/Ca ratios of Cibicides wuellerstorfi and Orbulina universa from Bahaman sediment cores

This geochemical investigation utilizes Ba/Ca in the benthic foraminifer Cibicides wuellerstorfi from cores taken from the Bahama Banks and the Caribbean Sea to reconstruct changes in basal thermocline ventilation (800-1000 m) and middepth thermohaline circulation (1000-2000 m) in the western North Atlantic during the last glacial period, focusing on the deglacial transition. Previous studies show that an increase in ventilation of the North Atlantic subtropical gyre during the Last Glacial Maximum (LGM) caused a 30-60% decrease in labile nutrients within the thermocline layer. Using foraminiferal Ba/Ca as a proxy of refractory nutrients, increased ventilation during the LGM produced a depletion of less than 20% compared to Holocene values. Following glaciation, the production of Glacial North Atlantic Intermediate Water (GNAIW) shut down owing to the presence of meltwater in the surface ocean, which resulted in a decrease in ventilation, as seen by an enrichment of barium in the basal thermocline. GNAIW was subsequently replaced by barium-rich southern component water in the middepth western North Atlantic. Foraminiferal Ba/Ca data suggest a 38% contribution from southern component water to a depth as shallow as 1475 m and a 14% contribution at 1123 m during deglaciation.

Benthic organic carbon flux and oxygen penetration depth in the Southern Ocean

For the investigation of organic carbon fluxes reaching the seafloor, oxygen microprofiles were measured at 145 sites in different sub-regions of the Southern Ocean. At eleven sites, an in situ oxygen microprofiler was deployed for the measurement of oxygen profiles and the calculation of organic carbon fluxes. At four sites, both in situ and ex situ data were determined for high latitudes. Based on this dataset as well as on previous published data, a relationship was established for the estimation of fluxes derived by ex situ measured O2 profiles. The fluxes of labile organic matter range from 0.5 to 37.1 mgC m**2/day. The high values determined by in situ measurements were observed in the Polar Front region (water depth of more than 4290 m) and are comparable to organic matter fluxes observed for high-productivity, upwelling areas like off West Africa. The oxygen penetration depth, which reflects the long-term organic matter flux to the sediment, was correlated with assemblages of key diatom species. In the Scotia Sea (~3000 m water depth), oxygen penetration depths of less than 15 cm were observed, indicating high benthic organic carbon fluxes. In contrast, the oxic zone extends down to several decimeters in abyssal sediments of the Weddell Sea and the southeastern South Atlantic. The regional pattern of organic carbon fluxes derived from micro-sensor data suggest that episodic and seasonal sedimentation pulses are important for the carbon supply to the seafloor of the deep Southern Ocean.

Viral, bacterial and ciliate abundance and viral-bacterial diversity in mesocosm experiments, 2007-2008, Kongsfjorden

Two mesocosm experiments, PAME-I and PAME-II were conducted in 2007 and 2008 to investigate fate of organic carbon in the arctic microbial food web. Mesocosms were nutrient fertilized initially to induce phytoplankton bloom development. In PAME-I eight units (each 700 L) formed two four point gradients of additional DOC in form of glucose (0, 0.5, 1 and 3 times Redfield ratio in terms of carbon relative to the nitrogen and phosphorus additions) (Fig. 1). All the eight units also got a daily dose of NH4+ and PO4**3- in Redfield ratio. Two gradients were set up, one with silicate addition, performed in the Arctic location Ny Ålesund, Svalbard, have previously been reported to give different food-web level responses to similar nutrient perturbations. In PAME-II all ten units (each 900 L) formed two four point gradients of additional DOC in form of glucose (0, 0.5, 1, 2 and 3 times Redfield ratio in terms of carbon relative to nitrogen and phosphorus additions). The two gradients in glucose were kept silicate replete. NH4+ was used as the DIN source in one gradient (units 1 to 5) and NO3- in the other (units 6-9). All units got a daily dose of PO4**3- in Redfield ratio. Prokaryotes and viruses were measured by flow cytometry, while ciliate abundances were counted using a Flow Cam. Viral and bacterial diversity was measured by PFGE and DGGE, respectively. In PAME-II the abundance of ciliates was lower than in PAME-I, presumably caused by higher copepod grazing. The abundances of prokaryotes and viruses were also lower in PAME-II compared to PAME-I. Further, less diversity was detected in the viral community (FCM and PFGE) in PAME-II, and no response was observed in the bacterial community structure due to addition of organic carbon.

(Table 3) Reduced inorganic sulphur, d34S and nickel content of ODP Leg 165 sites

Leg 165 of the Ocean Drilling Program afforded a unique opportunity to investigate organic and inorganic geochemistry across a wide gradient of sediment compositions and corresponding chemical pathways. The solid fractions at Sites 998, 999, 1000, and 1001 reveal varying proportions of reactive carbonate species, a labile volcanic ash fraction occurring in discrete layers and as a dispersed component, and detrital fluxes that derive from continental weathering. The relative proportions and reactivities of these end-members strongly dictate the character of the diagenetic profiles observed during the pore-water work of Leg 165. In addition, alteration of the well-characterized basaltic basement at Site 1001 has provided a strong signal that is reflected in many of the dissolved components. The relative effects of basement alteration and diagenesis within the sediment column are discussed in terms of downcore relationships for dissolved calcium and magnesium. With the exception of Site 1002 in the Cariaco Basin, the sediments encountered during Leg 165 were uniformly deficient in organic carbon (typically <0.1 wt%). Consequently, rates of organic oxidation were generally low and dominated by suboxic pathways with subordinate levels of bacterial sulfate reduction and methanogenesis. The low rates of organic remineralization are supported by modeled rates of sulfate reduction. Site 1000 provided an exception to the generally low levels of microbially mediated redox cycling. At this site the sediment is slightly more enriched in organic phases, and externally derived thermogenic hydrocarbons appear to aid in driving enhanced levels of redox diagenesis at great depths below the seafloor. The entrapment of these volatiles corresponds with a permeability seal defined by a pronounced Miocene minimum in calcium carbonate concentration recognized throughout the basin and with a dramatic downcore increase in the magnitude of limestone lithification. The latter has been tentatively linked to increases in alkalinity associated with microbial oxidation of organic matter and gaseous hydrocarbons. Recognition and quantification of previously unconstrained large volumes and frequencies of Eocene and Miocene silicic volcanic ash within the Caribbean Basin is one of the major findings of Leg 165. High frequencies of volcanic ash layers manifest as varied but often dominant controls on pore-water chemistry. Sulfur isotope results are presented that speak to secondary metal and sulfur enrichments observed in ash layers sampled during Leg 165. Ultimately, a better mechanistic understanding of these processes and the extent to which they have varied spatially and temporally may bear on the global mass balances for a range of major and minor dissolved components of seawater.

Extractions and analyses of reactive phosphorus in the Bering Sea from IODP Holes 323-U1341A and 323-U1341B

To reconstruct the cycling of reactive phosphorus (P) in the Bering Sea, a P speciation record covering the last ~ 4 Ma was generated from sediments recovered during Integrated Ocean Drilling Program (IODP) Expedition 323 at Site U1341 (Bowers Ridge). A chemical extraction procedure distinguishing between different operationally defined P fractions provides new insight into reactive P input, burial and diagenetic transformations. Reactive P mass accumulation rates (MARs) are ~ 20-110 µmol/cm2/ka, which is comparable to other open ocean locations but orders of magnitude lower than most upwelling settings. We find that authigenic carbonate fluorapatite (CFA) and opal-bound P are the dominant P fractions at Site U1341. An overall increasing contribution of CFA to total P with sediment depth is consistent with a gradual "sink switching" from more labile P fractions (fish remains, Fe oxides, organic matter) to stable authigenic CFA. However, the positive correlation of CFA with Al content implies that a significant portion of the supposedly reactive CFA is non-reactive "detrital contamination" by eolian and/or riverine CFA. In contrast to CFA, opal-bound P has rarely been studied in marine sediments. We find for the first time that opal-bound P directly correlates with excess silica contents. This P fraction was apparently available to biosiliceous phytoplankton at the time of sediment deposition and is a long-term sink for reactive P in the ocean, despite the likelihood for diagenetic re-mobilisation of this P at depth (indicated by increasing ratios of excess silica to opal-bound P). Average reactive P MARs at Site U1341 increase by ~ 25% if opal-bound P is accounted for, but decrease by ~ 25% if 50% of the extracted CFA fraction (based on the lowest CFA value at Site U1341) is assumed to be detrital. Combining our results with literature data, we present a qualitative perspective of terrestrial CFA and opal-bound P deposition in the modern ocean. Riverine CFA input has mostly been reported from continental shelves and margins draining P-rich lithologies, while eolian CFA input is found across wide ocean regions underlying the Northern Hemispheric "dust belt". Opal-bound P burial is important in the Southern Ocean, North Pacific, and likely in upwelling areas. Shifts in detrital CFA and opal-bound P deposition across ocean basins likely occurred over time, responding to changing weathering patterns, sea level, and biogenic opal deposition.

Cadmium analytical results from sediment and foraminifera shells

There has been a major contradiction between benthic foraminiferal Cd/Ca and d13C data concerning the labile nutrient chemistry of the Southern Ocean during the Last Glacial Maximum (LGM). Cd data indicates that LGM South Atlantic nutrient concentrations were as low as they are today, indicative of a persistent influx of nutrient-depleted North Atlantic Deep Water (NADW). d13C data indicates that LGM South Atlantic nutrient concentrations were much higher than at present (even higher than anywhere else in the ocean at that time), and these data have been interpreted as signifying the complete shutdown ofthe export of NADW into the global ocean. This paper examines both true geochemical differences and various confounding foraminiferal artifacts for both tracers. While many different processes and artifacts affect both tracers in the margin, we conclude the discrepancy is mainly due to the "Mackensen Effect" of low foraminiferal d13C as a result of high carbon flux to the sediments, and that LGM Atlantic Sector Southern Ocean nutrient concentrations remained similar to the levels encountered today.

Mean redox potential values in surface sediments along the transect D in the Cretan Sea (Table 1)

The seasonal, spatial and bathymetric changes in the distribution of chloroplastic pigments (Chl a, phaeopigments and CPE), TOC, TON, ATP, bottom water nutrient content and the main biochemical classes of organic compounds (lipids, proteins and carbohydrates) were recorded from May 1994 to September 1995 over the continental margin of northern Crete. The concentration of chloroplastic pigment equivalents (CPE) was always low, dropping dramatically along the shelf-slope gradient. Microbial activity (ATP) also dropped sharply beyond the continental shelf following a distribution pattern similar to TOC and TON. Lipid, protein and carbohydrate concentrations, as well as biopolymeric carbon were comparable to those reported for other more productive areas, however, the quality of the organic matter itself was rather poor. Thus, carbohydrates, the dominant biochemical class, were characterised by being highly (80-99%) refractory, as soluble carbohydrates represented (on annual average) only 6% of the total carbohydrate pool. Protein and lipid concentrations strongly decreased with depth, indicating depletion of trophic resources in the bathyal zone. Proteins appeared to be the more degradable compounds and indeed the protein to carbohydrate ratios were found to decrease strongly in the deeper stations. Organic matter content and quality decreased both with increasing distance from the coast and within the sediment. All sedimentary organic compounds were found to vary between sampling periods, with the changes being more pronounced over the continental shelf. The different temporal patterns of the various components suggest a different composition and/or origin of the OM inputs during the different sampling periods. The amount of material reaching the sediments below 540 m is extremely low, suggesting that most of the organic material is decomposed and/or utilised before reaching the sea floor. In conclusion, the continental shelf and bathyal sediments of the Cretan Sea can be considered, from a trophic point of view, as two different subsystems.

Meiofauna of surface sediments in the Cretan Sea

Quantitative information on metazoan meiofaunal abundance and biomass was obtained from three continental shelf (at 40, 100 and 200 m depth) and four deep-sea stations (at 540, 700, 940 and 1540 m depth) in the Cretan Sea (South Aegean Sea, NE Mediterranean). Samples were collected on a seasonal basis (from August 1994 to September 1995) with the use of a multiple corer. Meiofaunal abundance and biomass on the continental shelf of the Cretan Sea were high, in contrast to the extremely low values reported for the bathyal sediments that showed values comparable to those reported for abyssal and hadal environments. In order to explain the spatial and seasonal changes in metazoan meiofauna these data were compared with: (1) the concentrations of 'food indicators' (such as proteins, lipids, soluble carbohydrates and CPE) (2) the bacterial biomass (3) the flux of labile organic compounds to the sea floor at a fixed station (D7, 1540 m depth). Highly significant relationships between meiofaunal parameters and CPE, protein and lipid concentrations and bacterial biomass were found. Most of the indicators of food quality and quantity (such as CPE, proteins and carbohydrates) showed a clear seasonality with highest values in February and lowest in September. Such changes were more evident on the continental shelf rather than at deeper depths. On the continental shelf, significant seasonal changes in meiofaunal density were related to changes in the input of labile organic carbon whereas meiofaunal assemblages on the deep-sea stations showed time-lagged changes in response to the food input recorded in February 95. At all deep-sea stations meiofaunal density increased with a time lag of 2 months. Indications for a time-lagged meiofaunal response to the food inputs were also provided by the increase in nauplii densities during May 95 and the increase in individual biomass of nematodes, copepods and polychaetes between February and May 1995. The lack of strong seasonal changes in deep sea meiofaunal density suggests that the supply of organic matter below 500 m is not strong enough to support a significant meiofaunal development. Below 700 m depth >92% of the total biomass in the sediment was represented by bacteria. The ratio of bacterial to meiofaunal biomass increased with increasing water depth indicating that bacteria are probably more effective than meiofauna in exploiting refractory organic compounds. These data lead us to hypothesise that the deep-sea sediments of the Cretan Sea are largely dependent upon a benthic microbial loop.

Iron concentration and fluxes in the water column during POLARSTERN cruises in the Southern Ocean

In large parts of the Southern Ocean, primary production is limited due to shortage of iron (Fe). We measured vertical Fe profiles in the western Weddell Sea, Weddell-Scotia Confluence, and Antarctic Circumpolar Current (ACC), showing that Fe is derived from benthic Fe diffusion and sediment resuspension in areas characterized by high turbulence due to rugged bottom topography. Our data together with literature data reveal an exponential decrease of dissolved Fe (DFe) concentrations with increasing distance from the continental shelves of the Antarctic Peninsula and the western Weddell Sea. This decrease can be observed 3500 km eastward of the Antarctic Peninsula area, downstream the ACC. We estimated DFe summer fluxes into the upper mixed layer of the Atlantic sector of the Southern Ocean and found that horizontal advection dominates DFe supply, representing 54 ± 15% of the total flux, with significant vertical advection second most important at 29 ± 13%. Horizontal and vertical diffusion are weak with 1 ± 2% and 1 ± 1%, respectively. The atmospheric contribution is insignificant close to the Antarctic continent but increases to 15 ± 10% in the remotest waters (>1500 km offshore) of the ACC. Translating Southern Ocean carbon fixation by primary producers into biogenic Fe fixation shows a twofold excess of new DFe input close to the Antarctic continent and a one-third shortage in the open ocean. Fe recycling, with an estimated 'fe' ratio of 0.59, is the likely pathway to balance new DFe supply and Fe fixation.

Geochemistry and geomicrobiology profiles of ferruginous sediment, Lake Towuti, Indonesia

Lake Towuti is a tectonic basin, surrounded by ultramafic rocks. Lateritic soils form through weathering and deliver abundant iron (oxy)hydroxides but very little sulfate to the lake and its sediment. To characterize the sediment biogeochemistry, we collected cores at three sites with increasing water depth and decreasing bottom water oxygen concentrations. Microbial cell densities were highest at the shallow site - a feature we attribute to the availability of labile organic matter and the higher abundance of electron acceptors due to oxic bottom water conditions. At the two other sites, OM degradation and reduction processes below the oxycline led to partial electron acceptor depletion. Genetic information preserved in the sediment as extracellular DNA provides information on aerobic and anaerobic heterotrophs related to Actinobacteria, Nitrospirae, Chloroflexi and Thermoplasmatales. These taxa apparently played a significant role in the degradation of sinking organic matter. However, extracellular DNA concentrations rapidly decrease with core depth. Despite very low sulfate concentrations, sulfate-reducing bacteria were present and viable in sediments at all three sites, as confirmed by measurement of potential sulfate reduction rates. Microbial community fingerprinting supported the presence of taxa related to Deltaproteobacteria and Firmicutes with demonstrated capacity for iron and sulfate reduction. Concomitantly, sequences of Ruminococcaceae, Clostridiales and Methanomicrobiales indicated potential for fermentative hydrogen and methane production. Such first insights into ferruginous sediments show that microbial populations perform successive metabolisms related to sulfur, iron and methane. In theory, iron reduction could reoxidize reduced sulfur compounds and desorb OM from iron minerals to allow remineralization to methane. Overall, we found that biogeochemical processes in the sediments can be linked to redox differences in the bottom waters of the three sites, like oxidant concentrations and the supply of labile OM. At the scale of the lacustrine record, our geomicrobiological study should provide a means to link the extant subsurface biosphere to past environments.