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.

Grain entrainment and transport rates of magnetic minerals of coastal sediment samples from the Bay of Plenty, New Zealand

Heavy (magnetic & non-magnetic) minerals are found concentrated by natural processes in many fluvial, estuarine, coastal and shelf environments with a potential to form economic placer deposits. Understanding the processes of heavy mineral transport and enrichment is prerequisite to interpret sediment magnetic properties in terms of hydro- and sediment dynamics. In this study, we combine rock magnetic and sedimentological laboratory measurements with numerical 3D discrete element models to investigate differential grain entrainment and transport rates of magnetic minerals in a range of coastal environments (riverbed, mouth, estuary, beach and near-shore). We analyzed grain-size distributions of representative bulk samples and their magnetic mineral fractions to relate grain-size modes to respective transport modes (traction, saltation, suspension). Rock magnetic measurements showed that distribution shapes, population sizes and grain-size offsets of bulk and magnetic mineral fractions hold information on the transport conditions and enrichment process in each depositional environment. A downstream decrease in magnetite grain size and an increase in magnetite concentration was observed from riverine source to marine sink environments. Lower flow velocities permit differential settling of light and heavy mineral grains creating heavy mineral enriched zones in estuary settings, while lighter minerals are washed out further into the sea. Numerical model results showed that higher heavy mineral concentrations in the bed increased the erosion rate and enhancing heavy mineral enrichment. In beach environments where sediments contained light and heavy mineral grains of equivalent grain sizes, the bed was found to be more stable with negligible amount of erosion compared to other bed compositions. Heavy mineral transport rates calculated for four different bed compositions showed that increasing heavy mineral content in the bed decreased the transport rate. There is always a lag in transport between light and heavy minerals which increases with higher heavy mineral concentration in all tested bed compositions. The results of laboratory experiments were validated by numerical models and showed good agreement. We demonstrate that the presented approach bears the potential to investigate heavy mineral enrichment processes in a wide range of sedimentary settings.

Experimental evaluation of seaweeds as a vector for microplastics into marine food webs, supplementary material

The ingestion of microplastics has been shown for a great variety of marine organisms. However, benthic marine mesoherbivores such as the common periwinkle Littorina littorea have been largely disregarded in studies about the effects of microplastics on the marine biota, probably because the pathway for microplastics to this functional group of organisms was not obvious. In laboratory experiments we showed that the seaweed Fucus vesiculosus retains suspended microplastics on its surface. The numbers of microplastics that adhered to the algae correlated with the concentrations of suspended particles in the water. In choice feeding assays L. littorea did not distinguish between algae with adherent microplastics and clean algae without microplastics, indicating that the snails do not recognize solid nonfood particles in the submillimeter size range as deleterious. In periwinkles that were feeding on contaminated algae, microplastics were found in the stomach and in the gut. However, no microplastics were found in the midgut gland, which is the principle digestive organ of gastropods. Microplastics in the fecal pellets of the periwinkles indicate that the particles do not accumulate rapidly inside the animals but are mostly released with the feces. Our results provide the first evidence that seaweeds may represent an efficient pathway for microplastics from the water to marine benthic herbivores.

Calcium concentrations and isotopic composition in carbonates and porefluids from ODP Hole 130-807A

The calcium isotopic compositions (d44Ca) of 30 high-purity nannofossil ooze and chalk and 7 pore fluid samples from ODP Site 807A (Ontong Java Plateau) are used in conjunction with numerical models to determine the equilibrium calcium isotope fractionation factor (a_s-f) between calcite and dissolved Ca2+ and the rates of post-depositional recrystallization in deep sea carbonate ooze. The value of a_s-f at equilibrium in the marine sedimentary section is 1.0000+/-0.0001, which is significantly different from the value (0.9987+/-0.0002) found in laboratory experiments of calcite precipitation and in the formation of biogenic calcite in the surface ocean. We hypothesize that this fractionation factor is relevant to calcite precipitation in any system at equilibrium and that this equilibrium fractionation factor has implications for the mechanisms responsible for Ca isotope fractionation during calcite precipitation. We describe a steady state model that offers a unified framework for explaining Ca isotope fractionation across the observed precipitation rate range of ~14 orders of magnitude. The model attributes Ca isotope fractionation to the relative balance between the attachment and detachment fluxes at the calcite crystal surface. This model represents our hypothesis for the mechanism responsible for isotope fractionation during calcite precipitation. The Ca isotope data provide evidence that the bulk rate of calcite recrystallization in freshly-deposited carbonate ooze is 30-40%/Myr, and decreases with age to about 2%/Myr in 2-3 million year old sediment. The recrystallization rates determined from Ca isotopes for Pleistocene sediments are higher than those previously inferred from pore fluid Sr concentration and are consistent with rates derived for Late Pleistocene siliciclastic sediments using uranium isotopes. Combining our results for the equilibrium fractionation factor and recrystallization rates, we evaluate the effect of diagenesis on the Ca isotopic composition of marine carbonates at Site 807A. Since calcite precipitation rates in the sedimentary column are many orders of magnitude slower than laboratory experiments and the pore fluids are only slightly oversaturated with respect to calcite, the isotopic composition of diagenetic calcite is likely to reflect equilibrium precipitation. Accordingly, diagenesis produces a maximum shift in d44Ca of +0.15? for Site 807A sediments but will have a larger impact where sedimentation rates are low, seawater circulates through the sediment pile, or there are prolonged depositional hiatuses.

Tomographic measurements of O. umbonatus and N. truempyi during the PETM and ETM-2 at ODP Sites 208-1262 and 208-1263

Predicting the impact of ongoing anthropogenic CO2 emissions on calcifying marine organisms is complex, owing to the synergy between direct changes (acidification) and indirect changes through climate change (e.g., warming, changes in ocean circulation, and deoxygenation). Laboratory experiments, particularly on longer-lived organisms, tend to be too short to reveal the potential of organisms to acclimatize, adapt, or evolve and usually do not incorporate multiple stressors. We studied two examples of rapid carbon release in the geological record, Eocene Thermal Maximum 2 (~53.2 Ma) and the Paleocene Eocene Thermal Maximum (PETM, ~55.5 Ma), the best analogs over the last 65 Ma for future ocean acidification related to high atmospheric CO2 levels. We use benthic foraminifers, which suffered severe extinction during the PETM, as a model group. Using synchrotron radiation X-ray tomographic microscopy, we reconstruct the calcification response of survivor species and find, contrary to expectations, that calcification significantly increased during the PETM. In contrast, there was no significant response to the smaller Eocene Thermal Maximum 2, which was associated with a minor change in diversity only. These observations suggest that there is a response threshold for extinction and calcification response, while highlighting the utility of the geological record in helping constrain the sensitivity of biotic response to environmental change.

Photosynthetic production in the Central Arctic during POLARSTERN cruise ARK-XXVII/3 (IceArc) in 2012

The ice-covered Central Arctic Ocean is characterized by low primary productivity due to light and nutrient limitations. It has been speculated that the recent reduction in ice cover could lead to a substantial increase in primary production, but still little is known as to the fate of the ice-associated primary production, and of nutrient supply with increasing warming. This study presents results from the Central Arctic Ocean collected during summer 2012, when sea-ice reached a minimum extent since the onset of satellite observations. Net primary productivity (NPP) was measured in water column, sea ice and melt ponds by 14CO2 uptake at different irradiances. Photosynthesis vs. irradiance (PI) curves were established in laboratory experiments and used to upscale measured NPP to the deep Eurasian Basin (north of 78°N) using the irradiance-based Central Arctic Ocean Primary Productivity model (CAOPP). In addition, new annual production was calculated from the seasonal nutrient drawdown in the mixed layer since last winter. Results show that ice algae can contribute up to 60% to primary production in the Central Arctic at the end of the season. The ice-covered water column had lower NPP rates than open water probably due to light limitation. According to the nutrient ratios in the euphotic zone, nitrate limitation was detected in the Siberian Seas (Laptev Sea area), while silicate was the main limiting nutrient at the ice margin influenced by Atlantic waters. Although sea-ice cover was substantially reduced in 2012, total annual new production in the Eurasian Basin was 17 ± 7 Tg C/yr, which is similar to previous estimates. However, when including the contribution by sub-ice algal filaments, the annual production for the deep Eurasian Basin (north of 78°N) is 16 Tg C/yr higher than estimated before. Our data suggest that sub-ice algae might be responsible for potential local increases in NPP due to higher light availability under the ice, and their ability to benefit from a wider area of nutrients as they drift with the ice.

Radium concentration in water samples from the Northeast Atlantic

We present field measurements of air-sea gas exchange by the radon deficit method that were carried out during JASIN 1978 (NE Atlantic) and FGGE 1979 (Equatorial Atlantic). Both experiments comprised repeated deficit measurements at fixed position over periods of days or longer, using a previously descibed precise and fast-acquiaition, automatic radon measuring system. The deficit time series exhibit variations that only partly reflect the expected changes in gas transfer. By evaluating averages over each time series we deduce the following gas transfer velocities (average wind velocity and water temperature in parentheses): JASIN phase 1: 1.6 ± 0.8 m/d (at ~6 m/s, 13°C) JASIN phase 2: 4.3 ± 1.2 m/d (at ~8 m/s, 13°C) FGGE: 1.2 ± 0.4 m/d (at ~5 m/s, 28°C) 0.9 ± 0.4 m/d (at ~7 m/s, 28°C) 1.5 ± 0.4 m/d (at ~7 m/s, 28°C) The large difference betwen the JASIN phase 2 and FGGE values despite quite similare average wind velocity becomes even larger when the values are, however, fully compatible with the range of gas transfer velocities observed in laboratory experiments and the conclusion is suggested that their difference is caused by the highly different wind variability in JASIN and FGGE. We conclude that in gas exchange parameterization it is not sufficinent to consider wind velocity only. A comparison of our observations with laboratory results outlines the range of variations of air-sea gas transfer velocities with wind velocity and sea state. We also reformulate the radon deficit method, in the light of our observed deficit variations, to account explicitely for non-stationary and horizontal inhomogeneity in previous radon work introduces considerable uncertainty in deduced gas transfere velocity. We furthermore discuss the observational rewuirements that have to be met for an adequate exploitation of the radon deficit method, of which an observation area of minimum horizontal inhomogeneity and monitoring of the remaining inhomogeneities are thought to be the most stringent ones.

Data base on heterotrophic dinoflagellates grazing and growth rate as a function of size, prey size and type compiled from the literature

Microzooplankton (the 20 to 200 µm size class of zooplankton) is recognised as an important part of marine pelagic ecosystems. In terms of biomass and abundance heterotrophic dinoflagellates are one of the important groups of organism in microzooplankton. However, their rates - grazing and growth - , feeding behaviour and prey preferences are poorly known and understood. A set of data was assembled in order to derive a better understanding of heterotrophic dinoflagellates rates, in response to parameters such as prey concentration, prey type (size and species), temperature and their own size. With these objectives, literature was searched for laboratory experiments with information on one or more of these parameters effect studied. The criteria for selection and inclusion in the database included: (i) controlled laboratory experiment with a known dinoflagellate feeding on a known prey; (ii) presence of ancillary information about experimental conditions, used organisms - cell volume, cell dimensions, and carbon content. Rates and ancillary information were measured in units that meet the experimenter need, creating a need to harmonize the data units after collection. In addition different units can link to different mechanisms (carbon to nutritive quality of the prey, volume to size limits). As a result, grazing rates are thus available as pg C dinoflagellate-1 h-1, µm3 dinoflagellate-1 h-1 and prey cell dinoflagellate-1 h-1; clearance rate was calculated if not given and growth rate is expressed as the growth rate per day.

(Table 7) Representative electron microprobe analyses for natroalunite and larger FeOx spheroids of experiment ADSU6

Acid-sulfate alteration of basalt by SO2-bearing volcanic vapors has been proposed as one possible origin for sulfate-rich deposits on Mars. To better define mineralogical signatures of acid-sulfate alteration, laboratory experiments were performed to investigate alteration pathways and geochemical processes during reaction of basalt with sulfuric acid. Pyroclastic cinders composed of phenocrysts including plagioclase, olivine, and augite embedded in glass were reacted with sulfuric acid at 145 °C for up to 137 days at a range of fluid : rock ratios. During the experiments, the phenocrysts reacted rapidly to form secondary products, while the glass was unreactive. Major products included amorphous silica, anhydrite, and Fe-rich natroalunite, along with minor iron oxides/oxyhydroxides (probably hematite) and trace levels of other sulfates. At the lowest fluid : rock ratio, hexahydrite and an unidentified Fe-silicate phase also occurred as major products. Reaction-path models indicated that formation of the products required both slow dissolution of glass and kinetic inhibitions to precipitation of a number of minerals including phyllosilicates and other aluminosilicates as well as Al- and Fe-oxides/oxyhydroxides. Similar models performed for Martian basalt compositions predict that the initial stages of acid-sulfate alteration of pyroclastic deposits on Mars should result in formation of amorphous silica, anhydrite, Fe-bearing natroalunite, and kieserite, along with relict basaltic glass. In addition, analysis of the experimental products indicates that Fe-bearing natroalunite produces a Mössbauer spectrum closely resembling that of jarosite, suggesting that it should be considered an alternative to the component in sulfate-rich bedrocks at Meridiani Planum that has previously been identified as jarosite.

(Table 1) Compressional-wave velocities and anisotropies and wet-bulk densities of calcareous rocks and some cherts at DSDP Holes 62-463 and 62-465A

Anisotropy in compressional-wave velocities in sedimentary rocks recovered by DSDP has been recognized by several investigators (Boyce, 1976; Tucholke et al., 1976; Carlson and Christensen, 1977). The anisotropy is also observed at elevated pressures in laboratory experiments, and thus probably persists at depth in some calcareous rocks (Schreiber et al., 1972; Christensen et al., 1973; Carlson and Christensen, 1979). Carlson and Christensen (1979) suggested that the observed velocity anisotropy was produced not by the alignment of cracks but by the alignment of c axes of calcite perpendicular to bedding during compaction, diagenesis, and recrystallization. On DSDP Leg 62, calcareous rocks were recovered from the western Mid-Pacific Mountains (sub-bottom depths of 452-823 m, Site 463) and southern Hess Rise (276-412 m, Site 465). Most of the calcareous rocks are horizontally laminated and color-banded, and ages are early Cenomanian to late Barremian (Site 463 and 465 reports, this volume). The purpose of this study is to confirm the velocity anisotropy in the calcareous rocks and to identify any relationship of anistropy to bulk density, mean velocity, and burial depth.

Fecal pellet and egg production rates of copepod species at varying suspended sediment concentrations

We investigated the effect of suspended sediments on the vital rates of the copepods Calanus finmarchicus, Pseudocalanus sp. and Metridia longa in a Greenland sub-Arctic fjord. The fjord had a gradient of suspended particulate matter (SPM) with high concentrations (>50 mg/L) in the inner fjord due to glacial melt water runoff. Laboratory experiments showed that when feeding on the diatom Thalassiosira weissflogii specific ingestion rates were low at high concentrations of suspended sediment for C. finmarchicus (>20 mg/L) and Pseudocalanus sp. (>50 mg/L), while no effect was found for M. longa. For C. finmarchicus, a relatively constant fecal pellet production (FPP) and fecal pellet volume suggested ingestion of sediment, which probably led to reduction in egg production rates (EPRs) at high sediment concentrations. For Pseudocalanus sp., FPP decreased with increasing sediment concentrations, while no effect was observed on EPR. No significant difference was observed in FPP for M. longa feeding on the diatom T. weissflogii compared to the ciliate Strombidium sulcatum. The study shows that high sediment concentrations influence the capability of carbon turnover in C. finmarchicus and Pseudocalanus sp., while M. longa appears to be more tolerant to high sediment loads. Therefore, high concentrations of SPM could potentially influence the species composition of glacially influenced fjords.

Marine meso-herbivore consumption scales faster with temperature than seaweed primary production, supplementary material

Respiration of ectotherms is predicted to increase faster with rising environmental temperature than photosynthesis of primary producers because of the differential temperature dependent kinetics of the key enzymes involved. Accordingly, if biological processes at higher levels of complexity are constrained by underlying metabolic functions food consumption by heterotrophs should increase more rapidly with rising temperature than photo-autoptrophic primary production. We compared rates of photosynthesis and growth of the benthic seaweed Fucus vesiculosus with respiration and consumption of the isopod Idotea baltica to achieve a mechanistic understanding why warming strengthens marine plant-herbivore interactions. In laboratory experiments thallus pieces of the seaweed and individuals of the grazer were exposed to constant temperatures at a range from 10 to 20°C. Photosynthesis of F. vesiculosus did not vary with temperature indicating efficient thermal acclimation whereas growth of the algae clearly increased with temperature. Respiration and food consumption of I. baltica also increased with temperature. Grazer consumption scaled about 2.5 times faster with temperature than seaweed production. The resulting mismatch between algal production and herbivore consumption may result in a net loss of algal tissue at elevated temperatures. Our study provides an explanation for faster decomposition of seaweeds at elevated temperatures despite the positive effects of high temperatures on algal growth.

Geochemistry of pore water and sediments offshore Nicaragua and Costa Rica

Four volcanic ash-bearing marine sediment cores and one ash-free reference core were examined during research cruise RV Meteor 54/2 offshore Nicaragua and Costa Rica to investigate the chemical composition of pore waters related to volcanic ash alteration. Sediments were composed of terrigenous matter derived from the adjacent continent and contained several distinct ash layers. Biogenic opal and carbonate were only minor components. The terrigenous fraction was mainly composed of smectite and other clay minerals while the pore water composition was strongly affected by the anaerobic degradation of particulate organic matter via microbial sulphate reduction. The alteration of volcanic matter showed only a minor effect on major element concentrations in pore waters. This is in contrast to prior studies based on long sediment cores taken during the DSDP, where deep sediments always showed distinct signs of volcanic ash alteration. The missing signal of ash alteration is probably caused by low reaction rates and the high background concentration of major dissolved ions in the seawater-derived pore fluids. Dissolved silica concentrations were, however, significantly enriched in ash-bearing cores and showed no relation to the low but variable contents of biogenic opal. Hence, the data suggest that silica concentrations were enhanced by ash dissolution. Thus, the dissolved silica profile measured in one of the sediment cores was used to derive the in-situ dissolution rate of volcanic glass particles in marine sediments. A non-steady state model was run over a period of 43 kyr applying a constant pH of 7.30 and a dissolved Al concentration of 0.05 ?M. The kinetic constant (AA) was varied systematically to fit the model to the measured dissolved silica-depth profile. The best fit to the data was obtained applying AA = 1.3 * 10**-U9 mol of Si/cm**2/ s. This in-situ rate of ash dissolution at the seafloor is three orders of magnitude smaller than the rate of ash dissolution determined in previous laboratory experiments. Our results therefore imply that field investigations are necessary to accurately predict natural dissolution rates of volcanic glasses in marine sediments.

Permeability measurements on sheared and unsheared samples from different IODP Holes of Leg 316 and from IODP Hole 315-C0001E and ODP Hole 190-1174B

At subduction zones, the permeability of major fault zones influences pore pressure generation, controls fluid flow pathways and rates, and affects fault slip behavior and mechanical strength by mediating effective normal stress. Therefore, there is a need for detailed and systematic permeability measurements of natural materials from fault systems, particularly measurements that allow direct comparison between the permeability of sheared and unsheared samples from the same host rock or sediment. We conducted laboratory experiments to compare the permeability of sheared and uniaxially consolidated (unsheared) marine sediments sampled during IODP Expedition 316 and ODP Leg 190 to the Nankai Trough offshore Japan. These samples were retrieved from: (1) The décollement zone and incoming trench fill offshore Shikoku Island (the Muroto transect); (2) Slope sediments sampled offshore SW Honshu (the Kumano transect) ~ 25 km landward of the trench, including material overriden by a major out-of-sequence thrust fault, termed the "megasplay"; and (3) A region of diffuse thrust faulting near the toe of the accretionary prism along the Kumano transect. Our results show that shearing reduces fault-normal permeability by up to 1 order of magnitude, and this reduction is largest for shallow (< 500 mbsf) samples. Shearing-induced permeability reduction is smaller in samples from greater depth, where pre-existing fabric from compaction and lithification may be better developed. Our results indicate that localized shearing in fault zones should result in heterogeneous permeability in the uppermost few kilometers in accretionary prisms, which favors both the trapping of fluids beneath and within major faults, and the channeling of flow parallel to fault structure. These low permeabilities promote the development of elevated pore fluid pressures during accretion and underthrusting, and will also facilitate dynamic hydrologic processes within shear zones including dilatancy hardening and thermal pressurization.

Organic carbon and humic substances contents, carbon isotope composition and peroxidase activity in sediments from DSDP Sites 15-147 and 75-532

Sediment samples from the Cariaco Trench (DSDP Leg 15) and the Walvis Ridge (DSDP Leg 75) ranging in age from Holocene to Upper Miocene (approximately 8 million years BP) and in depth from 5 to 258 m were extracted with basic sodium pyrophosphate and the extract analyzed for enzymic activity. Since no dehydrogenase, alkaline phosphatase or esterase activity was found, it is estimated from these data that the maximum bacterial population does not exceed 1000 cells per gram dry sediment. Peroxidase activity was, however, found in most samples: this showed marked dependence on the humic substance concentration (expressed as percent of the organic carbon content) and increased with depth at a rate of 33 units per meter. To explain this observation, we favor an hypothesis based on the presence of active humic-enzyme association. The humic substances absorb and stabilize peroxidase which is liberated throughout the sediment column by lysis of cells. The association of the enzyme with the humic substances protects it from biodegradation and denaturation. This hypothesis agrees with laboratory experiments which show the enhanced stability of humic-enzyme complexes towards degradation by biological, chemical and thermal effects.