Data from PhD thesis

The ocean quahog, Arctica islandica is the longest-lived non-colonial animal known to science. A maximum individual age of this bivalve of 405 years has been found in a population off the north western coast of Iceland. Conspicuously shorter maximum lifespan potentials (MLSPs) were recorded from other populations of A. islandica in European waters (e.g. Kiel Bay: 30 years, German Bight: 150 years) which experience wider temperature and salinity fluctuations than the clams from Iceland. The aim of my thesis was to identify possible life-prolonging physiological strategies in A. islandica and to examine the modulating effects of extrinsic factors (e.g. seawater temperature, food availability) and intrinsic factors (e.g. species-specific behavior) on these strategies. Burrowing behavior and metabolic rate depression (MRD), tissue-specific antioxidant and anaerobic capacities as well as cell-turnover (= apoptosis and proliferation) rates were investigated in A. islandica from Iceland and the German Bight. An inter-species comparison of the quahog with the epibenthic scallop Aequipecten opercularis (MLSP = 8-10 years) was carried out in order to determine whether bivalves with short lifespans and different lifestyles also feature a different pattern in cellular maintenance and repair. The combined effects of a low-metabolic lifestyle, low oxidative damage accumulation, and constant investment into cellular protection and tissue maintenance, appear to slow-down the process of physiological aging in A. islandica and to afford the extraordinarily long MLSP in this species. Standard metabolic rates were lower in A. islandica when compared to the shorter-lived A. opercularis. Furthermore, A. islandica regulate mantle cavity water PO2 to mean values < 5 kPa, a PO2 at which the formation of reactive oxygen species (ROS) in isolated gill tissues of the clams was found to be 10 times lower than at normoxic conditions (21 kPa). Burrowing and metabolic rate depression (MRD) in Icelandic specimens were more pronounced in winter, possibly supported by low seawater temperature and food availability, and seem to be key energy-saving and life-prolonging parameters in A. islandica. The signaling molecule nitric oxide (NO) may play an important role during the onset of MRD in the ocean quahog by directly inhibiting cytochome-c-oxidase at low internal oxygenation upon shell closure. In laboratory experiments, respiration of isolated A. islandica gills was completely inhibited by chemically produced NO at low experimental PO2 <= 10 kPa. During shell closure, mantle cavity water PO2 decreased to 0 kPa for longer than 24 h, a state in which ROS production is supposed to subside. Compared to other mollusk species, onset of anaerobic metabolism is late in A. islandica in the metabolically reduced state. Increased accumulation of the anaerobic metabolite succinate was initially detected in the adductor muscle of the clams after 3.5 days under anoxic incubation or in burrowed specimens. A ROS-burst was absent in isolated gill tissue of the clams following hypoxia (5 kPa)-reoxygenation (21 kPa). Accordingly, neither the activity of antioxidant enzymes superoxide dismutase (SOD) and catalase (CAT), nor the specific content of the ROS-scavenger glutathione (GSH) was enhanced in different tissues of the ocean quahog after 3.5 days of self-induced or forced hypoxia/anoxia to prepare for an oxidative burst. While reduced ROS formation compared to routine levels lowers oxidative stress during MRD and also during surfacing, the general preservation of high cellular defense and the efficient removal and replacement of damaged cells over lifetime seem to be of crucial importance in decelerating the senescent decline in tissues of A. islandica. Along with stable antioxidant protection over 200 years of age, proliferation rates and apoptosis intensities in most investigated tissues of the ocean quahog were low, but constant over 140 years of age. Accordingly, age-dependent accumulations of protein and lipid oxidation products are lower in A. islandica tissues when compared to the shorter-lived bivalve A. opercularis. The short-lived swimming scallop is a model bivalve species representing the opposite life and aging strategy to A. islandica. In this species permanently high energy throughput, reduced investment into antioxidant defense with age, and higher accumulation of oxidation products are met by higher cell turnover rates than in the ocean quahog. The only symptoms of physiological change over age ever found in A. islandica were decreasing cell turnover rates in the heart muscle over a lifetime of 140 years. This may either indicate higher damage levels and possibly ongoing loss of functioning in the heart of aging clams, or, the opposite, lower rates of cell damage and a reduced need for cell renewal in the heart tissue of A. islandica over lifetime. Basic physiological capacities of different A. islandica populations, measured at controlled laboratory conditions, could not explain considerable discrepancies in population specific MLSPs. For example, levels of tissue-specific antioxidant capacities and cell turnover rates were similarly high in individuals from the German Bight and from Iceland. Rather than genetic differences, the local impacts of environmental conditions on behavioral and physiological traits in the ocean quahog seem to be responsible for differences in population-specific MLSPs.

Composition of recent carbonate bottom sediments from the Guiana shelf an off the West Africa

This work was based on a study of the upper layer of recent carbonate bottom sediments of the Atlantic Ocean. Biogenic carbonate of recent sediments is represented by metastable and stable minerals. In the ocean metastable phases can exist indefinitely long, but the structure of polymorphism determines inevitability of transformation of metastable phases into stable ones. This transformation occurs in the solid phase. In the absence of a critical point between the two phases of the transition process is not available for study by microscopic methods. It is estimated indirectly by studying the nature and extent of changes in mineral and chemical compositions. With aging of sediments their mineral composition alters in direction of increasing contents of resistant minerals. Fine grained sediments and fractions are subject to more intensive effects of early diagenesis processes, rather than coarse ones; this is reflected in their mineral composition. Regularities of distribution of carbonate minerals in size fractions consistent with the direction of polymorphic transformations in calcium carbonate. Such transformations can occur in a particular dimension of grains. Concrete grain size depends on environmental conditions. This situation explains presence of metastable biogenic carbonates at different depths of the ocean and suggests presence of diagenetic calcite in sediments occurring below expected for each case depth of the transition.

Radiocarbon ages and plateau definitions of sediments from the abyssal ocean

Ice core records demonstrate a glacial-interglacial atmospheric CO2 increase by ~100 ppm, while 14C calibration efforts document a strong decrease in atmospheric 14C concentration during this period. A calculated transfer of ~530 Gt of 14C depleted carbon is required to produce the deglacial coeval rise of carbon in the atmosphere and terrestrial biosphere. This amount is usually ascribed to oceanic carbon release, although the actual mechanisms remained elusive, since an adequately old and carbon-enriched deep-ocean reservoir seemed unlikely. Here we present a new, though still fragmentary, ocean-wide d14C dataset showing that during the Last Glacial Maximum (LGM) and Heinrich Stadial 1 (HS-1) the maximum 14C age difference between ocean deep waters and the atmosphere exceeded the modern values by up to 1500 14C yr, in the extreme reaching 5100 14C yr. Below 2000 m depth the 14C ventilation age of modern ocean waters is directly linked to the concentration of dissolved inorganic carbon (DIC). We propose as working hypothesis that the modern regression of DIC vs d14C also applies for LGM times, which implies that a mean LGM aging by ~600 14C yr corresponded to a global rise of ~85-115 µmol DIC/kg in the deep ocean. Thus, the prolonged residence time of ocean deep waters may indeed have made it possible to absorb an additional ~730-980 Gt DIC, one third of which possibly originated from intermediate waters. We also infer that LGM deep-water O2 dropped to suboxic values of <10µmol/kg in the Atlantic sector of the Southern Ocean, possibly also in the subpolar North Pacific. The outlined deglacial transfer of the extra aged, deep-ocean carbon to the atmosphere via the dynamic ocean-atmosphere carbon exchange would be sufficient to account for two trends observed, (1) for the increase in atmospheric CO2 and (2) for the 190-permil drop in atmospheric d14C during the so-called HS-1 'Mystery Interval', when atmospheric 14C production rates were largely constant.

Mineral and oxygen isotopic compositions in a profile through the altered lower oceanic crust, ODP Hole 176-735B

ODP Hole 735B located on the Southwest Indian Ridge at 57°E is an in situ sampled long, continuous section of lower oceanic crust. Oxygen isotope compositions of constituent minerals of Leg 176 gabbros have been measured by UV-laser oxygen isotope microprobe. Together with existing data from Leg 118, a complete oxygen isotope profile through the lower oceanic crust has been obtained. Most clinopyroxenes and olivines have normal mantle values of ~5.5 per mil and ~5.2 per mil, respectively, while plagioclases show slight d18O enrichment relative to its mantle value of 6.1per mil. Down-hole variations of Hole 735B gabbro indicate a downward decreasing d18O profile, with a kink at a depth of about 800 m below sea floor. Above this depth, gabbros are depleted in 18O relative to unaltered basalts, while below ~800 m they show nearly unmodified d18O values. Abundant seawater penetration appears to be limited to the upper part of the lower crust at ODP site 735 (~800 m into the gabbroic layer and ~2-2.5 km into the oceanic crust from the top of pillow basalts). Mass balance calculations show that the lower crust formed under this ultra-slow-spreading ridge has an average d18O value of 5.5 per mil. The whole crust at Site 735 has an overall 18O enrichment with d18O values of 6.0 per mil to 7.8 per mil, depending on the possible variation of the d18O values of the upper pillow basalts and sheeted dykes. The apparent difference in oxygen isotope compositions of ocean crusts formed with different spreading rates has important implications on the buffering of ocean water over geological time, as well as on the oxygen recycling between crust and mantle through subduction. The difference of seawater penetration between fast- and slow-spreading ridges could be related to their particular magmatic-tectonic history during the formation and aging of the crust. However, more analyses on continuous sections through oceanic and ophiolitic crust in different tectonic settings are required to derive any predictive models.

Geochemical composition and atomic formulas of saponites, celadonites and carbonates from ODP Leg 168 sites

With this study, we investigate the mineralogical variations associated with the low-temperature (<100°C) alteration of normal tholeiitic pillow basalts varying in age from 0.8 to 3.5 Ma. Their alteration intensity varies systematically and is related to several factors, including (1) the aging of the igneous crust, (2) the increase of temperatures from the younger to the older sites, measured at the sediment/basement interface, (3) the local and regional variations in lithology and primary porosity, and (4) the degree of pillow fracturing. Fractures represent the most important pathways that allow significant penetration of fluids into the rock and are virtually the only factor controlling the alteration of the glassy rim and the early stages of pillow alteration. Three different alteration stages have been recognized: alteration of glassy margin, oxidizing alteration through fluid circulation in fracture systems, and reducing alteration through diffusion. All the observed mineralogical and chemical variations occurring during the early stages of alteration are interpreted as the result of the rock interaction with "normal," alkaline, and oxidizing seawater, along preferential pathways represented by the concentric and radial crack systems. The chemical composition of the fluid progressively evolves while moving into the basalt, leading to a reducing alteration stage, which is initially responsible for the precipitation of Fe-rich saponite and minor sulfides and subsequently for the widespread formation of carbonates. At the same time, the system evolved from being "water dominated" to being "rock dominated." No alteration effects in pillow basalts were observed that must have occurred at temperatures higher than those measured during Leg 168 at the basement/sediment interface (e.g., between 15° and 64°C).

Working on a baseline for the Kongsfjorden food web: production and properties of macroalgal particulate organic matter (POM), supplementary material

Macroalgae, in particular kelps, produce a large amount of biomass in Kongsfjorden, which is to a great extent released into the water in an annual cycle. As an example, the brown alga Alaria esculenta loses its blade gradually, 3 ± 0.8 % of the blade area per day (August 2012), thereby adding to the pool of particulate organic matter (POM) in the fjord. Upon release small thallus pieces are "aging" in that they are prone to leaching and serving as substrate for microorganisms, thus turning into palatable food for suspension and bottom feeders. In order to define a macroalgal baseline for the Kongsfjorden food web, stable isotopes d14C and d15N were measured in individuals of A. esculenta, Saccharina latissima and Laminaria digitata directly sampled after collection and in artificially produced POM (aPOM) of A. esculenta that was allowed to age under experimental conditions. In aPOM from this species sampled in August 2012 the C/N ratios decreased between d1 and d8 of a 14-day culture period in parallel to the fading photosynthetic activity of the algal fragments as demonstrated by use of an Imaging-PAM. Microscopic observations of the aPOM in August 2012 and 2013 revealed the frequent occurrence of small brown algal endo- and epiphytes. First feeding experiments with Mysis oculata (Mysids) and Hiatella arctica (Bivalves) showed that these species can ingest macroalgal POM. The importance of kelp-derived POM for the food web is subject of the current research.

Chemistry of low-temperature hydrothermal altered basement of ODP Site 129-801

Low-temperature hydrothermal alteration of basement from Site 801 was studied through analyses of the mineralogy, chemistry, and oxygen isotopic compositions of the rocks. The more than 100-m section of 170-Ma basement consists of 60 m of tholeiitic basalt separated from the overlying 60 m of alkalic basalts by a >3-m-thick Fe-Si hydrothermal deposit. Four alteration types were distinguished in the basalts: (1) saponite-type (Mg-smectite) rocks are generally slightly altered, exhibiting small increases in H2O, d18O, and oxidation; (2) celadonite-type rocks are also slightly altered, but exhibit uptake of alkalis in addition to hydration and oxidation, reflecting somewhat greater seawater/rock ratios than the saponite type; (3) Al-saponite-type alteration resulted in oxidation, hydration, and alkali and 18O uptake and losses of Ca and Na due to the breakdown of plagioclase and clinopyroxene; and (4) blue-green rocks exhibit the greatest chemical changes, including oxidation, hydration, alkali uptake, and loss of Ca, Na, and Mg due to the complete breakdown of plagioclase and olivine to K-feldspar and phyllosilicates. Saponite- and celadonite-type alteration of the tholeiite section occurred at a normal mid-ocean ridge basalt spreading center at temperatures <20°C. Near- or off-axis intrusion of an alkali basalt magma at depth reinitiated hydrothermal circulation, and the Fe-Si hydrothermal deposit formed from cool (<60°C) distal hydrothermal fluids. Focusing of fluid flow in the rocks immediately underlying the deposit resulted in the extensive alteration of the blue-green rocks at similar temperatures. Al-saponite alteration of the subsequent alkali basalts overlying the deposit occurred at relatively high water/rock ratios as part of the same low-temperature circulation system that formed the hydrothermal deposit. Abundant calcite formed in the rocks during progressive "aging" of the crust during its long history away from the spreading center.

Stable oxygen and carbon isotope record of benthic foraminifera in Pliocene sediments of the Californian margin

New benthic foraminiferal stable isotopic records of northeast Pacific intermediate water (ODP Site 1014, 1177 m) and mid-depth water (ODP Site 1018, 2476 m) were compared to isotopic records of deep water in the tropical Pacific (ODP Site 849, 3851 m) for the reconstruction of vertical profiles of nutrient and physical properties from the Early Pliocene to the Early Pleistocene (approx. 5-1.4 Ma). Our data indicate that, for the entire interval, there was enhanced north Pacific intermediate water ventilation relative to today, and a similar to modern circulation pattern with northward flowing Pacific Bottom Water (PBW) beneath its southward flowing return flow. However, the core of maximally aged return flow resided as deep as ~2500 m (as compared to ~1500 m today), probably due to the strengthened intermediate water flow. Less apparent aging of deep water along its path before 2.7 Ma indicates that thermohaline overturning may have been more rapid in the warm period of the Early Pliocene. In addition, prior to 2.7 Ma, foraminiferal oxygen isotopic values at mid-depth sites are higher than at deep sites (a reversed vertical gradient relative to today) in both the Atlantic and Pacific Oceans. We suggest that NADW was warmer and more saline than today and that it influenced mid-depth waters throughout the Atlantic and Pacific Oceans. Enhanced Pliocene formation of warmer/saltier intermediate water in the north Pacific, and deep water in the north Atlantic, may have been a result of higher than modern high/mid-latitude sea surface temperatures, evaporation, and salinity.

Stable isotope ratios of benthic foraminifera in mid-Pleitsocene sediments of ODP Site 108-664 in the equatorial Atlantic

Five delta13C records from the deep ocean, extending back to 1.3 Ma, were examined in order to constrain changes in mean ocean carbon isotope composition and thermohaline circulation over the 41- to 100-ka climate transition. These data show that significant perturbations in mean ocean carbon chemistry were associated with the mid-Pleistocene climate transition. Notable features of the last 1.3 Myr are (1) a pronounced ~0.3? decrease in mean ocean delta13C between 0.9 and 1.0 Myr, followed by a return to pre-1.0 Ma values by 400 ka B.P., which we propose was due to the onetime addition of isotopically depleted terrestrial carbon to the ocean, possibly associated with an increase in global aridity (and decrease in the size of the biosphere) across the 41- to 100-ka transition; (2) no change in the Atlantic-Pacific (A-P) delta13C gradient over the last 1.3 Myr, suggesting no change in mean ocean nutrient content accompanied the addition of light carbon; and (3) stronger vertical nutrient fractionation in the North Atlantic in the middle Pleistocene between sites 607 and 552, suggesting weaker North Atlantic Deep Water formation at this time relative to the early and late Pleistocene. We also find evidence for a more pronounced deep recirculation gyre in the western North Atlantic basin in the early Brunhes, as evidenced by "aging" of deep northern basin water (site 607) relative to deep water in the equatorial Atlantic (site 664).

Ostracoda in five sediment cores from the Laptev and Kara seas

Fossil ostracods were investigated in five AMS14C-dated cores from different parts of the Laptev and Kara seas. Three cores from the Laptev Sea shelf are located in river paleovalleys, and one core originates from the western continental slope. The core from the Kara Sea was obtained in the eastern shelf region. Six fossil assemblages were distinguished: estuarine (1), inner-shelf (2), middle-shelf (3), outer-shelf (4), Pre-Holocene upper continental slope (5), and Holocene upper continental slope (6). They show that during the Postglacial sea-level rise there was a transition from estuarine brackish-water environment to modern marine conditions. Assemblages 1-3 are present in the eastern Laptev Sea with the oldest ostracod-bearing samples aging back to 11.4-11.3 cal.ka. Cores from the western Laptev Sea (12.3 cal.ka, assemblages 1-4) and the Kara Sea (8.1 cal.ka, assemblages 2-4) demonstrate similar pattern in assemblage replacement, but contain a number of relatively deep-water species reflecting stronger influence of open-sea waters. Core from the continental slope, water depth 270 m (~ 17 cal.ka) encompasses assemblages 5 and 6, which are absent in the shelf cores. Assemblage 5 stands out as a specific community dominated by relatively deep-water Arctic and North Atlantic species together with euryhaline ones. The assemblages indicate inflows of Atlantic-derived waters and downslope slides due to the proximity to the paleocoastline. Assemblage (6) is similar to the modern local ostracod assemblage at this site.

Chlorofluorocarbons and noble gas measured on water bottle samples during three POLARSTERN cruises

We use a 27 year long time series of repeated transient tracer observations to investigate the evolution of the ventilation time scales and the related content of anthropogenic carbon (Cant) in deep and bottom water in the Weddell Sea. This time series consists of chlorofluorocarbon (CFC) observations from 1984 to 2008 together with first combined CFC and sulphur hexafluoride (SF6) measurements from 2010/2011 along the Prime Meridian in the Antarctic Ocean and across the Weddell Sea. Applying the Transit Time Distribution (TTD) method we find that all deep water masses in the Weddell Sea have been continually growing older and getting less ventilated during the last 27 years. The decline of the ventilation rate of Weddell Sea Bottom Water (WSBW) and Weddell Sea Deep Water (WSDW) along the Prime Meridian is in the order of 15-21%; the Warm Deep Water (WDW) ventilation rate declined much faster by 33%. About 88-94% of the age increase in WSBW near its source regions (1.8-2.4 years per year) is explained by the age increase of WDW (4.5 years per year). As a consequence of the aging, the Cant increase in the deep and bottom water formed in the Weddell Sea slowed down by 14-21% over the period of observations.

Chemistry of serpentinized peridotites from ODP Hole 109-670A

In this paper we describe textural relationships in hydrated upper mantle peridotites emplaced at a nonconstructive ridge segment. Development of serpentinites and partially serpentinized peridotites takes place in four main stages: (1) pervasive serpentinization forming mainly lizardite, (2) a tensional stage forming chrysotile + talc + chlorite, (3) a deformational stage forming antigorite + tremolite, and (4) a late local tensional stage forming another generation of chrysotile veinlets. Mineral chemistry of serpentine pseudomorphs reflects in part primary mineral compositions. Olivine pseudomorphs are typically nickeliferous and depleted in aluminum and chromium. Orthopyroxene pseudomorphs have lower nickel contents and relatively high iron, aluminum, and chromium contents. Clinopyroxene pseudomorphs have very low nickel contents and relatively high aluminum and chromium contents. These chemical patterns in the serpentinites can be used to help discriminate between harzburgitic and lherzolitic protoliths. Oxygen isotopes and mineral parageneses suggest serpentine is derived from circulation of hydrothermal (200?C) fluids through the peridotite body. Crystallization of tremolite, talc, and chlorite may have occurred at temperatures up to 525?C if C02/H20 ratios were less than 0.25. Open fissures developing in aging upper mantle provide paths for important seawater circulation through a thin basaltic carapace down to shallow mantle rocks.

Marine fungi from the Northwest African upwelling areas and from the Atlantic off Portugal

Quantitative data on lower marine Phycomycetes (fungi) found in the upwelling waters off the West African coast during cruises No. 13 (1968), 19 (1970), 36 (1975) and 44 (1977) of R.V. "Meteor" are reported. The distribution of the total fungi numbers is presented and, as far as possible, the evaluation of the material up to species level is given. Several provisionally named forms and groups of morphologically related, undescribed fungi are included. A correlation between the number of fungi in sediments and the water depth and distance from the coast line is postulated. There are typical distributions of the lower marine fungi in water bodies and sediments. Different values within replicates of the stations in different years show that there is a sequence in development of fungal populations induced by changes in the water bodies. Surface water far from the coast has low numbers of fungi; numbers increase to a maximum nearer to the coast. In the vicinity of the coast the values decrease. The numbers of fungi in the deep sediments are low below 1,200 m. However, there are isolated areas of higher fungal activities, indicated by some deeper grab samples. During two cruises, the "overlying water" in the grab samples was investigated. It was evident that the numbers of fungi lost by stirring of the sediment when the grab was brought up to the surface were small, relatively and absolutely. The seamount "Josephine Bank" has been investigated for the first time with respect to lower marine fungi; the populations are low in the sediments, but one sample of the surface water had a higher number than the water in the surroundings. In some hydrographic series there was a peculiar depth distribution. An increase occurred at a depth greater than 1,000 m. The results are discussed and some correlations to the aging of the fungal populations in the water masses are constructed.

(Table 1) Calcium carbonate and <2 µm fractions of bulk sample and insoluble residue at DSDP Hole 92-597

Particles of red brown to yellow brown semiopaque oxides (RSO) dominate the insoluble residue fraction of the sediments at Site 597. Unlike the X-ray amorphous particles in the Bauer Deep sediments, these particles are composed of mainly goethite; the amount of X-ray amorphous ferric hydroxide and poorly crystalline ferromanganese oxyhydroxides is generally small relative to the amount of goethite. A qualitative goethite crystallinity index was established. The variations observed in the crystallinity of goethite with increasing depth and changes in lithology suggest that aging and long-term exposure to seawater in a high water/sediment regime influence and increase the rate of recrystallization of the Fe-oxyhydroxides of the RSO particles. The percentage of organic carbon is low in these sediments; it varies primarily between 0.2 and 0.4 wt.%. Phillipsite is present throughout the sediment column and is more concentrated in the youngest clay layer and in the oldest basal sediments.