Heat stress effects on crop performance and tools for tolerance breeding

Abiotic stress is one of the most common causes of crop deficit and loss and hence an important area of study. Moreover, concerns regarding global climate change over past decades mean the study of different abiotic stresses appears to be essential if its effects are to be mitigated. The current review covers the effects of heat stress on crop performance, the response crops make when subjected to this stress and the development of tools designed to breed for stress tolerant crops. Distinct levels of the problem are considered, from the morphological/anatomical, through the physiological and to the biochemical/molecular. The study of heat shock proteins (HSPs), quantitative trait loci (QTLs) identification and the relationship between metabolomics (OMICS) and heat stress are given special consideration.

Stable isotope analysis of benthic foraminifera and sedimentology on ODP Site 162-984

Sediments recovered at ODP Site 984 on the Reykjanes Ridge provided multicentennial-scale records (SST, planktic and benthic delta18O, IRD and magnetic susceptibility) of Late Pliocene climate change over the onset of Northern Hemisphere glaciation (NHG), 2.95-2.82 Ma. Short-term climate variations prior and after the onset of continent-wide glaciation were compared to test the hypothesis, whether Dansgaard-Oeschger (DO) cycles may have been triggered by continental ice breakouts. By means of spectral analyses for two selected interglacial stages prior to and after NHG (G15 and G1), we found that climate variability resembled that of the Holocene and the mid-Pliocene warm period. In contrast, DO-like periodicities near 1470, 2900, and 4400 yr indeed only occurred in glacial stages after the onset of NHG (G14, G6, and 104), but hardly in stage G20 prior to the onset. These results suggest a causal link between DO cycles and the Late Pliocene onset of major NHG and ice breakouts in the North Atlantic. This data set provides all primary data and spectral analysis related to this scientific work.

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.

Lipid, carbon and nitrogen content and fatty acid composition of Temora longicornis females and different food species during the experiment

Temora longicornis, a dominant calanoid copepod species in the North Sea, is characterised by low lipid reserves and high biomass turnover rates. To survive and reproduce successfully, this species needs continuous food supply and thus requires a highly flexible digestive system to exploit various food sources. Information on the capacity of digestive enzymes is scarce and therefore the aim of our study was to investigate the enzymatic capability to respond to quickly changing nutritional conditions. We conducted two feeding experiments with female T. longicornis from the southern North Sea off Helgoland. In the first experiment in 2005, we tested how digestive enzyme activities and enzyme patterns as revealed by substrate SDS-PAGE (sodium dodecylsulfate polyacrylamide gel electrophoresis) responded to changes in food composition. Females were incubated for three days fed ad libitum with either the heterotrophic dinoflagellate Oxyrrhis marina or the diatom Thalassiosira weissflogii. At the beginning and at the end of the experiment, copepods were deep-frozen for analyses. The lipolytic enzyme activity did not change over the course of the experiment but the enzyme patterns did, indicating a distinct diet-induced response. In a second experiment in 2008, we therefore focused on the enzyme patterns, testing how fast changes occur and whether feeding on the same algal species leads to similar patterns. In this experiment, we kept the females for 4 days at surplus food while changing the algal food species daily. At day 1, copepods were offered O. marina. On day 2, females received the cryptophycean Rhodomonas baltica followed by T. weissflogii on day 3. On day 4 copepods were again fed with O. marina. Each day, copepods were frozen for analysis by means of substrate SDS-PAGE. This showed that within 24 h new digestive enzymes appeared on the electrophoresis gels while others disappeared with the introduction of a new food species, and that the patterns were similar on day 1 and 4, when females were fed with O. marina. In addition, we monitored the fatty acid compositions of the copepods, and this indicated that specific algal fatty acids were quickly incorporated. With such short time lags between substrate availability and enzyme response, T. longicornis can successfully exploit short-term food sources and is thus well adapted to changes in food availability, as they often occur in its natural environment due seasonal variations in phyto- and microzooplankton distribution.

Grain size composition and age of ODP Hole 167-1017E sediments

Siliciclastic sedimentation at Ocean Drilling Program Site 1017 on the southern slope of the Santa Lucia Bank, central California margin, responded closely to oceanographic and climatic change over the past ~130 ka. Variation in mean grain-size and sediment sorting within the ~25-m-thick succession from Hole 1017E show Milankovitch-band to submillenial-scale variation. Mean grain size of the "sortable silt" fraction (10-63 µm) ranges from 17.6 to 33.9 µm (average 24.8 µm) and is inversely correlated with the degree of sorting. Much of the sediment has a bimodal or trimodal grain-size distribution that is composed of distinct fine silt, coarse silt to fine sand, and clay-size components. The position of the mode and the sorting of each component changes through the succession, but the primary variation is in the presence or abundance of the coarse silt fraction that controls the overall mean grain size and sorting of the sample. The occurrence of the best-sorted, finest grained sediment at high stands of sea level (Holocene, marine isotope Substages 5c and 5e) reflect the linkage between global climate and the sedimentary record at Site 1017 and suggest that the efficiency of off-shelf transport is a key control of sedimentation on the Santa Lucia Slope. It is not clear what proportion of the variation in grain size and sorting may also be caused by variations in bottom current strength and in situ hydrodynamic sorting.

Comparisons of observed ice properties and Envisat Advanced Synthetic Aperture Radar (ASAR) data during Nathanial B. Palmer cruise NBP09-01 and Oden cruise OSO08/09

Envisat Advanced Synthetic Aperture Radar (ASAR) Wide Swath Mode (WSM) images are used to derive C-band HH-polarization normalized radar cross sections (NRCS). These are compared with ice-core analysis and visual ship-based observations of snow and ice properties observed according to the Antarctic Sea Ice Processes and Climate (ASPeCt) protocol during two International Polar Year summer cruises (Oden 2008 and Palmer 2009) in West Antarctica. Thick first-year (TFY) and multi-year (MY) ice were the dominant ice types. The NRCS value ranges between -16.3 ± 1.1 and -7.6 ± 1.0 dB for TFY ice, and is -12.6 ± 1.3 dB for MY ice; for TFY ice, NRCS values increase from ~-15 dB to -9 dB from December/January to mid-February. In situ and ASPeCt observations are not, however, detailed enough to interpret the observed NRCS change over time. Co-located Advanced Microwave Scanning Radiometer-Earth Observing System (AMSR-E) vertically polarized 37 GHz brightness temperatures (TB37V), 7 day and 1 day averages as well as the TB37V difference between ascending and descending AMSR-E overpasses suggest the low NRCS values (-15 dB) are associated with snowmelt being still in progress, while the change towards higher NRCS values (-9dB) is caused by commencement of melt-refreeze cycles after about mid-January.

Late Quaternary carbonate and isotope stratigraphy of DSDP Site 90-594 east of South Island, New Zealand

The late Quaternary sequence off eastern South Island, New Zealand, consists of ~100 m of alternating bluish gray pelagic oozes and greenish gray hemipelagic oozes that extend uninterruptedly back to the Brunhes/Matuyama boundary (0.73 m.y.). A very high resolution (~2400 yr.) record of sediment texture, calcium carbonate content, and planktonic and benthic foraminiferal oxygen and carbon isotope composition demonstrates an in-phase cyclical fluctuation between the sedimentary parameters that closely correspond to the pelagic-hemipelagic sedimentation cycles and the isotope composition. Pelagic oozes, formed during interglacial periods of high eustatic sea level, are characterized by calcareous microfossils, relative enrichment in sand and clay sizes, high carbonate contents, reduced delta18O values, and increased delta13C values. Hemipelagic oozes, associated with glacial episodes and lowered eustatic sea level, include common terrigenous material and siliceous microfossils, are enriched in silt sizes, have low carbonate contents, high delta18O values, and low delta13C values. The history of alpine glaciations and associated erosion of the South Island of New Zealand, as expressed by the appearance of hemipelagic oozes, can be correlated directly with the major fluctuations of Northern Hemisphere ice sheets as expressed by the influence of eustatic sea-level changes on the oxygen isotope composition of both planktonic and benthic foraminifers. This high-accumulation-rate record contains conspicuous intervals of highfrequency, high-amplitude isotope variability including the presence of multiple glacial/interglacial intervals within single isotope stages, and offers one of the best sections cored to date for detailed study of the evolution and history of climate change over the last 0.75 m.y.

Sea ice thickness from airborne electromagnetic (EM) induction sounding acquired from fixed-wing aircraft POLAR 5 surveys from several airports in Spitsbergen, Greenland, Canada and Alaska

While summer Arctic sea-ice extent has decreased over the past three decades, it is subject to large interannual and regional variations. Methodological challenges in measuring ice thickness continue to hamper our understanding of the response of the ice-thickness distribution to recent change, limiting the ability to forecast sea-ice change over the next decade. We present results from a 2400 km long pan-Arctic airborne electromagnetic (EM) ice thickness survey in April 2009, the first-ever large-scale EM thickness dataset obtained by fixed-wing aircraft over key regions of old ice in the Arctic Ocean between Svalbard and Alaska. The data provide detailed insight into ice thickness distributions characteristic for the different regions. Comparison with previous EM surveys shows that modal thicknesses of old ice had changed little since 2007, and remained within the expected range of natural variability.

(Table 1) Movement parameters of nine adult female snowy owls (Bubo scandiacus) tracked during the winter period 07/08 and 08/09 around Hudson Strait, Canada

Top predators of the arctic tundra are facing a long period of very low prey availability during winter and subsidies from other ecosystems such as the marine environment may help to support their populations. Satellite tracking of snowy owls, a top predator of the tundra, revealed that most adult females breeding in the Canadian Arctic overwinter at high latitudes in the eastern Arctic and spend several weeks (up to 101 d) on the sea-ice between December and April. Analysis of high-resolution satellite images of sea-ice indicated that owls were primarily gathering around open water patches in the ice, which are commonly used by wintering seabirds, a potential prey. Such extensive use of sea-ice by a tundra predator considered a small mammal specialist was unexpected, and suggests that marine resources subsidize snowy owl populations in winter. As sea-ice regimes in winter are expected to change over the next decades due to climate warming, this may affect the wintering strategy of this top predator and ultimately the functioning of the tundra ecosystem.

Shifts in the microbial community structure in different temperatures during sample storage from IODP Hole 325-M0058A

The objective of this study was to determine shifts in the microbial community structure and potential function based on standard Integrated Ocean Drilling Program (IODP) storage procedures for sediment cores. Standard long-term storage protocols maintain sediment temperature at 4°C for mineralogy, geochemical, and/or geotechnical analysis whereas standard microbiological sampling immediately preserves sediments at -80°C. Storage at 4°C does not take into account populations may remain active over geologic time scales at temperatures similar to storage conditions. Identification of active populations within the stored core would suggest geochemical and geophysical conditions within the core change over time. To test this potential, the metabolically active fraction of the total microbial community was characterized from IODP Expedition 325 Great Barrier Reef sediment cores prior to and following a 3-month storage period. Total RNA was extracted from complementary 2, 20, and 40 m below sea floor sediment samples, reverse transcribed to complementary DNA and then sequenced using 454 FLX sequencing technology, yielding over 14,800 sequences from the six samples. Interestingly, 97.3% of the sequences detected were associated with lineages that changed in detection frequency during the storage period including key biogeochemically relevant lineages associated with nitrogen, iron, and sulfur cycling. These lineages have the potential to permanently alter the physical and chemical characteristics of the sediment promoting misleading conclusions about the in situ biogeochemical environment. In addition, the detection of new lineages after storage increases the potential for a wider range of viable lineages within the subsurface that may be underestimated during standard community characterizations.

Stable isotope record, opal accumulation rate and geochemistry of middle Miocene sediments of IODP Site 321-U1338

During the middle Miocene, Earth's climate transitioned from a relatively warm phase (Miocene climatic optimum) into a colder mode with re-establishment of permanent ice sheets on Antarctica, thus marking a fundamental step in Cenozoic cooling. Carbon sequestration and atmospheric CO2 drawdown through increased terrestrial and/or marine productivity have been proposed as the main drivers of this fundamental transition. We integrate high-resolution (1-3 k.y.) benthic stable isotope data with XRF-scanner derived biogenic silica and carbonate accumulation estimates in an exceptionally well-preserved sedimentary archive, recovered at Integrated Ocean Drilling Program Site U1338, to reconstruct eastern equatorial Pacific productivity variations and to investigate temporal linkages between high- and low-latitude climate change over the interval 16-13 Ma. Our records show that the climatic optimum (16.8-14.7 Ma) was characterized by high amplitude climate variations, marked by intense perturbations of the carbon cycle. Episodes of peak warmth at (southern hemisphere) insolation maxima coincided with transient shoaling of the carbonate compensation depth and enhanced carbonate dissolution in the deep ocean. A switch to obliquity-paced climate variability after 14.7 Ma concurred with a general improvement in carbonate preservation and the onset of stepwise global cooling, culminating with extensive ice growth over Antarctica at ~13.8 Ma. We find that two massive increases in opal accumulation at ~14.0 and ~13.8 Ma occurred just before and during the final and most prominent cooling step, supporting the hypothesis that enhanced siliceous productivity in the eastern equatorial Pacific contributed to CO2 drawdown.

Age determination and stable isotope record of ODP Hole 162-980B

The conversion of surface water to deep water in the North Atlantic results in the release of heat from the ocean to the atmosphere, which may have amplified millennial-scale climate variability during glacial times (Broecker et al., 1990, doi:10.1029/PA005i004p00469) and could even have contributed to the past 11,700 years of relatively mild climate (known as the Holocene epoch) (Bond et al., 2001, doi:10.1126/science.1065680; Alley et al., 1997, doi:10.1130/0091-7613(1997)025<0483:HCIAPW>2.3.CO;2; Keigwin and Boyle, 2000, doi:10.1073/pnas.97.4.1343). Here we investigate changes in the carbon-isotope composition of benthic foraminifera throughout the Holocene and find that deep-water production varied on a centennial-millennial timescale. These variations may be linked to surface and atmospheric events that hint at a contribution to climate change over this period.

(Table 1) Water storage changes of the 33 world's largest river basins between 2002-2009

Global change in land water storage and its effect on sea level is estimated over a 7-year time span (August 2002 to July 2009) using space gravimetry data from GRACE. The 33 World largest river basins are considered. We focus on the year-to-year variability and construct a total land water storage time series that we further express in equivalent sea level time series. The short-term trend in total water storage adjusted over this 7-year time span is positive and amounts to 80.6 ± 15.7 km**3/yr (net water storage excess). Most of the positive contribution arises from the Amazon and Siberian basins (Lena and Yenisei), followed by the Zambezi, Orinoco and Ob basins. The largest negative contributions (water deficit) come from the Mississippi, Ganges, Brahmaputra, Aral, Euphrates, Indus and Parana. Expressed in terms of equivalent sea level, total water volume change over 2002-2009 leads to a small negative contribution to sea level of -0.22 ± 0.05 mm/yr. The time series for each basin clearly show that year-to-year variability dominates so that the value estimated in this study cannot be considered as representative of a long-term trend. We also compare the interannual variability of total land water storage (removing the mean trend over the studied time span) with interannual variability in sea level (corrected for thermal expansion). A correlation of ~0.6 is found. Phasing, in particular, is correct. Thus, at least part of the interannual variability of the global mean sea level can be attributed to land water storage fluctuations.

Geochemical data derived from an absolutely dated shell-based chronology for the Irish Sea

The identification in various proxy records of periods of rapid (decadal scale) climate change over recent millennia, together with the possibility that feedback mechanisms may amplify climate system responses to increasing atmospheric CO2, highlights the importance of a detailed understanding, at high spatial and temporal resolutions, of forcings and feedbacks within the system. Such an understanding has hitherto been limited because the temperate marine environment has lacked an absolute timescale of the kind provided by tree-rings for the terrestrial environment and by corals for the tropical marine environment. Here we present the first annually resolved, multi-centennial (489-year), absolutely dated, shell-based marine master chronology. The chronology has been constructed by detrending and averaging annual growth increment widths in the shells of multiple specimens of the very long-lived bivalve mollusc Arctica islandica, collected from sites to the south and west of the Isle of Man in the Irish Sea. The strength of the common environmental signal expressed in the chronology is fully comparable with equivalent statistics for tree-ring chronologies. Analysis of the 14C signal in the shells shows no trend in the marine radiocarbon reservoir correction (DR), although it may be more variable before ~1750. The d13C signal shows a very significant (R**2 = 0.456, p < 0.0001) trend due to the 13C Suess effect.