Preconditioning in the reef-building coral Pocillopora damicornis and the potential for trans-generational acclimatization in coral larvae under future climate change conditions

Coral reefs are globally threatened by climate change-related ocean warming and ocean acidification (OA). To date, slow-response mechanisms such as genetic adaptation have been considered the major determinant of coral reef persistence, with little consideration of rapid-response acclimatization mechanisms. These rapid mechanisms such as parental effects that can contribute to trans-generational acclimatization (e.g. epigenetics) have, however, been identified as important contributors to offspring response in other systems. We present the first evidence of parental effects in a cross-generational exposure to temperature and OA in reef-building corals. Here, we exposed adults to high (28.9°C, 805 µatm PCO2) or ambient (26.5°C, 417 µatm PCO2) temperature and OA treatments during the larval brooding period. Exposure to high treatment negatively affected adult performance, but their larvae exhibited size differences and metabolic acclimation when subsequently re-exposed, unlike larvae from parents exposed to ambient conditions. Understanding the innate capacity corals possess to respond to current and future climatic conditions is essential to reef protection and maintenance. Our results identify that parental effects may have an important role through (1) ameliorating the effects of stress through preconditioning and adaptive plasticity, and/or (2) amplifying the negative parental response through latent effects on future life stages. Whether the consequences of parental effects and the potential for trans-generational acclimatization are beneficial or maladaptive, our work identifies a critical need to expand currently proposed climate change outcomes for corals to further assess rapid response mechanisms that include non-genetic inheritance through parental contributions and classical epigenetic mechanisms.

Seawater carbonate chemistry and swimming behaviors of the raphidophyte Heterosigma akashiwo in a laboratory experiment

We investigated the effects of pH on movement behaviors of the harmful algal bloom causing raphidophyte Heterosigma akashiwo. Motility parameters from >8000 swimming tracks of individual cells were quantified using 3D digital video analysis over a 6-h period in 3 pH treatments reflecting marine carbonate chemistry during the pre-industrial era, currently, and the year 2100. Movement behaviors were investigated in two different acclimation-to-target-pH conditions: instantaneous exposure and acclimation of cells for at least 11 generations. There was no negative impairment of cell motility when exposed to elevated PCO2 (i.e., low pH) conditions but there were significant behavioral responses. Irrespective of acclimation condition, lower pH significantly increased downward velocity and frequency of downward swimming cells (p < 0.001). Rapid exposure to lower pH resulted in 9% faster downward vertical velocity and up to 19% more cells swimming downwards (p < 0.001). Compared to pH-shock experiments, pre-acclimation of cells to target pH resulted in ~30% faster swimming speed and up to 46% faster downward velocities (all p < 0.001). The effect of year 2100 PCO2 levels on population diffusivity in pre-acclimated cultures was >2-fold greater than in pH-shock treatments (2.2 × 105 µm**2/s vs. 8.4 × 104 µm**2/s). Predictions from an advection-diffusion model, suggest that as PCO2 increased the fraction of the population aggregated at the surface declined, and moved deeper in the water column. Enhanced downward swimming of H. akashiwo at low pH suggests that these behavioral responses to elevated PCO2 could reduce the likelihood of dense surface slick formation of H. akashiwo through reductions in light exposure or growth independent surface aggregations. We hypothesize that the HAB alga's response to higher PCO2 may exploit the signaling function of high PCO2 as indicative of net heterotrophy in the system, thus indicative of high predation rates or depletion of nutrients.

Physiology, growth and development of larval krill ''Euphausia superba'' in autumn and winter in the Lazarev Sea, Antarctica

The physiological condition of larval Antarctic krill was investigated during austral autumn 2004 and winter 2006 in the Lazarev Sea, to provide better understanding of a critical period of their life cycle. The condition of larvae was quantified in both seasons by determining their body length (BL), dry mass (DM), elemental- and biochemical composition, as well as stomach content analysis, and rates of metabolism and growth. Overall the larvae in autumn were in better condition under the ice than in open water, and for those under the ice there was a decrease in condition from autumn to winter. Thus growth rates of furcilia larvae in open water in autumn were similar to winter values under the ice (mean 0.008 mm/d), whereas autumn, under ice values were higher: 0.015 mm/d. Equivalent larval stages had up to 30% lower BL and 70% lower DM in winter compared to autumn, with mean oxygen consumption 44% lower (0.54 µl O2 DM/h). However, their ammonium excretion rates doubled (from 0.03-0.06 µg NH4 DM/h) so their mean O:N ratio was 46 in autumn and 15 in winter. Thus differing metabolic substrates were used between autumn and winter, suggesting a flexible overwintering strategy, as suggested for adults. The larvae were eating small copepods (Oithona spp.) and/or protozoans as well as autotrophic food under the ice. However, pelagic Chlorophyll a (Chl a) was a good predictor for growth in both seasons. The physics (current speed/ice topography) probably has a critical part to play in whether larval krill can exploit the food that may be associated with sea ice or be advected away from such suitable feeding habitat.

Depth profiles of dissolved sulfide and sulfate in the pore water of hydrocarbon seep sediments offshore Pakistan

The interaction between fluid seepage, bottom water redox, and chemosynthetic communities was studied at cold seeps across one of the world's largest oxygen minimum zones (OMZ) located at the Makran convergent continental margin. Push cores were obtained from seeps within and below the core-OMZ with a remotely operated vehicle. Extracted sediment pore water was analyzed for sulfide and sulfate concentrations. Depending on oxygen availability in the bottom water, seeps were either colonized by microbial mats or by mats and macrofauna. The latter, including ampharetid polychaetes and vesicomyid clams, occurred in distinct benthic habitats, which were arranged in a concentric fashion around gas orifices. At most sites colonized by microbial mats, hydrogen sulfide was exported into the bottom water. Where macrofauna was widely abundant, hydrogen sulfide was retained within the sediment. Numerical modeling of pore water profiles was performed in order to assess rates of fluid advection and bioirrigation. While the magnitude of upward fluid flow decreased from 11 cm yr**-1 to <1 cm yr**-1 and the sulfate/methane transition (SMT) deepened with increasing distance from the central gas orifice, the fluxes of sulfate into the SMT did not significantly differ (6.6-9.3 mol m**-2 yr**-1). Depth-integrated rates of bioirrigation increased from 120 cm yr**-1 in the central habitat, characterized by microbial mats and sparse macrofauna, to 297 cm yr**-1 in the habitat of large and few small vesicomyid clams. These results reveal that chemosynthetic macrofauna inhabiting the outer seep habitats below the core-OMZ efficiently bioirrigate and thus transport sulfate down into the upper 10 to 15 cm of the sediment. In this way the animals deal with the lower upward flux of methane in outer habitats by stimulating rates of anaerobic oxidation of methane (AOM) with sulfate high enough to provide hydrogen sulfide for chemosynthesis. Through bioirrigation, macrofauna engineer their geochemical environment and fuel upward sulfide flux via AOM. Furthermore, due to the introduction of oxygenated bottom water into the sediment via bioirrigation, the depth of the sulfide sink gradually deepens towards outer habitats. We therefore suggest that - in addition to the oxygen levels in the water column, which determine whether macrofaunal communities can develop or not - it is the depth of the SMT and thus of sulfide production that determines which chemosynthetic communities are able to exploit the sulfide at depth. We hypothesize that large vesicomyid clams, by efficiently expanding the sulfate zone down into the sediment, could cut off smaller or less mobile organisms, as e.g. small clams and sulfur bacteria, from the sulfide source.

PIES time series in the North Atlantic at the Mid-Atlantic Ridge (2006-2010)

In the western North Atlantic, warm and saline water is brought by the North Atlantic Current (NAC) from the subtropics into the subpolar gyre. Four inverted echo sounders with high precision pressure sensors (PIES) were moored between 47°40' N and 52°30' N to study the main pathways of the NAC from the western into the eastern basin. The array configuration that forms three segments (northern, central, and southern) allows partitioning of the NAC and some assessment of NAC flow paths through the different Mid-Atlantic Ridge fracture zones. We exploit the correlation between the NAC transport measured between 2006 and 2010 and the geostrophic velocity from altimeter data to extend the time series of NAC transports to the period from 1992 to 2013. The mean NAC transport over the entire 21 years is 27 ± 5 Sv, consisting of 60% warm water of subtropical origin and 40% subpolar water. We did not find a significant trend in the total transport time series, but individual segments had opposing trends, leading to a more focused NAC in the central subsection and decreasing transports in the southern and northern segments. The spectral analysis exhibits several significant peaks. The two most prominent are around 120 days, identified as the time scale of meanders and eddies, and at 4-9 years, most likely related to the NAO. Transport composites for the years of highest and lowest NAO indices showed a significantly higher transport (+2.9 Sv) during strong NAO years, mainly in the southern segment.

Age tie points and geochemistry of sediment cores from the Antarctic continental margin

Few high-latitude terrestrial records document the timing and nature of the Cenozoic "Greenhouse" to "Icehouse" transition. Here we exploit the bulk geochemistry of marine siliciclastic sediments from drill cores on Antarctica's continental margin to extract a unique semiquantitative temperature and precipitation record for Eocene to mid-Miocene (~54-13 Ma). Alkaline elements are strongly enriched in the detrital mineral fraction in fine-grained siliciclastic marine sediments and only occur as trace metals in the biogenic fraction. Hence, terrestrial climofunctions similar to the chemical index of alteration (CIA) can be applied to the alkaline major element geochemistry of marine sediments on continental margins in order to reconstruct changes in precipitation and temperature. We validate this approach by comparison with published paleotemperature and precipitation records derived from fossil wood, leaves, and pollen and find remarkable agreement, despite uncertainties in the calibrations of the different proxies. A long-term cooling on the order of >=8°C is observed between the Early Eocene Climatic Optimum (~54-52 Ma) and the middle Miocene (~15-13 Ma) with the onset of transient cooling episodes in the middle Eocene at ~46-45 Ma. High-latitude stratigraphic records currently exhibit insufficient temporal resolution to reconstruct continental aridity and inferred ice-sheet development during the middle to late Eocene (~45-37 Ma). However, we find an abrupt aridification of East Antarctica near the Eocene-Oligocene transition (~34 Ma), which suggests that ice coverage influenced high-latitude atmospheric circulation patterns through albedo effects from the earliest Oligocene onward.

Benthic megafaunal composition at the Arctic deep-sea observatory HAUSGARTEN

The Long-Term Ecological Research (LTER) observatory HAUSGARTEN, in the eastern Fram Strait, provides us the valuable ability to study the composition of benthic megafaunal communities through the analysis of seafloor photographs. This, in combination with extensive sampling campaigns, which have yielded a unique data set on faunal, bacterial, biogeochemical and geological properties, as well as on hydrography and sedimentation patterns, allows us to address the question of why variations in megafaunal community structure and species distribution exist within regional (60-110 km) and local (<4 km) scales. Here, we present first results from the latitudinal HAUSGARTEN gradient, consisting of three different stations (N3, HG-IV, S3) between 78°30'N and 79°45'N (2351 - 2788 m depth), obtained via the analysis of images acquired by a towed camera (OFOS - Ocean Floor Observation System) in 2011. We assess variability in megafaunal densities, species composition and diversity as well as biotic and biogenic habitat features, which may cause the patterns observed. While there were significant regional-scale differences in megafaunal composition and densities between the stations (N3 = 26.74 ± 0.63; HG-IV = 11.21 ± 0.25; S3 = 18.34 ± 0.39 individuals/m**2), significant local differences were only found at HG-IV. Regional-scale variations may be due to the significant differences in ice coverage at each station as well as the different quantities of protein available, whereas local-scale differences at HG-IV may be a result of variation in bottom topography or factors not yet identified.

Nitrogen fuelling of the pelagic food web of the Tropical Atlantic

We estimated the relative contribution of atmospheric Nitrogen (N) input (wet and dry deposition and N fixation) to the epipelagic food web by measuring N isotopes of different functional groups of epipelagic zooplankton along 23°W (17°N-4°S) and 18°N (20-24°W) in the Eastern Tropical Atlantic. Results were related to water column observations of nutrient distribution and vertical diffusive flux as well as colony abundance of Trichodesmium obtained with an Underwater Vision Profiler (UVP5). The thickness and depth of the nitracline and phosphocline proved to be significant predictors of zooplankton stable N isotope values. Atmospheric N input was highest (61% of total N) in the strongly stratified and oligotrophic region between 3 and 7°N, which featured very high depth-integrated Trichodesmium abundance (up to 9.4×104 colonies m-2), strong thermohaline stratification and low zooplankton delta15N (~2 per mil). Relative atmospheric N input was lowest south of the equatorial upwelling between 3 and 5°S (27%). Values in the Guinea Dome region and north of Cape Verde ranged between 45 and 50%, respectively. The microstructure-derived estimate of the vertical diffusive N flux in the equatorial region was about one order of magnitude higher than in any other area (approximately 8 mmol m-2 d 1). At the same time, this region received considerable atmospheric N input (35% of total). In general, zooplankton delta15N and Trichodesmium abundance were closely correlated, indicating that N fixation is the major source of atmospheric N input. Although Trichodesmium is not the only N fixing organism, its abundance can be used with high confidence to estimate the relative atmospheric N input in the tropical Atlantic (r2 = 0.95). Estimates of absolute N fixation rates are two- to tenfold higher than incubation-derived rates reported for the same regions. Our approach integrates over large spatial and temporal scales and also quantifies fixed N released as dissolved inorganic and organic N. In a global analysis, it may thus help to close the gap in oceanic N budgets.