Impact of thermal stress on evolutionary trajectories of pathogen resistance in three-spined stickleback (Gasterosteus aculeatus), supplementary material

Background: Pathogens are a major regulatory force for host populations, especially under stressful conditions. Elevated temperatures may enhance the development of pathogens, increase the number of transmission stages, and can negatively influence host susceptibility depending on host thermal tolerance. As a net result, this can lead to a higher prevalence of epidemics during summer months. These conditions also apply to marine ecosystems, where possible ecological impacts and the population-specific potential for evolutionary responses to changing environments and increasing disease prevalence are, however, less known. Therefore, we investigated the influence of thermal stress on the evolutionary trajectories of disease resistance in three marine populations of three-spined sticklebacks Gasterosteus aculeatus by combining the effects of elevated temperature and infection with a bacterial strain of Vibrio sp. using a common garden experiment. Results: We found that thermal stress had an impact on fish weight and especially on survival after infection after only short periods of thermal acclimation. Environmental stress reduced genetic differentiation (QST) between populations by releasing cryptic within-population variation. While life history traits displayed positive genetic correlations across environments with relatively weak genotype by environment interactions (GxE), environmental stress led to negative genetic correlations across environments in pathogen resistance. This reversal of genetic effects governing resistance is probably attributable to changing environment-dependent virulence mechanisms of the pathogen interacting differently with host genotypes, i.e. GPathogenxGHostxE or (GPathogenxE)x(GHostxE) interactions, rather than to pure host genetic effects, i.e. GHostxE interactions. Conclusion: To cope with climatic changes and the associated increase in pathogen virulence, host species require wide thermal tolerances and pathogen-resistant genotypes. The higher resistance we found for some families at elevated temperatures showed that there is evolutionary potential for resistance to Vibrio sp. in both thermal environments. The negative genetic correlation of pathogen resistance between thermal environments, on the other hand, indicates that adaptation to current conditions can be a weak predictor for performance in changing environments. The observed feedback on selective gradients exerted on life history traits may exacerbate this effect, as it can also modify the response to selection for other vital components of fitness.

Theoretical model simulations for variations in thermal reaction norms

Thermal reaction norms for growth rates of six Emiliania huxleyi isolates originating from the central Atlantic (Azores, Portugal) and five isolates from the coastal North Atlantic (Bergen, Norway) were assessed. We used the template mode of variation model to decompose variations in growth rates into modes of biological interest: vertical shift, horizontal shift, and generalist-specialist variation. In line with the actual habitat conditions, isolates from Bergen (Bergen population) grew well at lower temperatures, and isolates from the Azores (Azores population) performed better at higher temperatures. The optimum growth temperature of the Azores population was significantly higher than that of the Bergen population. Neutral genetic differentiation was found between populations by microsatellite analysis. These findings indicate that E. huxleyi populations are adapted to local temperature regimes. Next to between-population variation, we also found variation within populations. Genotype-by-environment interactions resulted in the most pronounced phenotypic differences when isolates were exposed to temperatures outside the range they naturally encounter. Variation in thermal reaction norms between and within populations emphasizes the importance of using more than one isolate when studying the consequences of global change on marine phytoplankton. Phenotypic plasticity and standing genetic variation will be important in determining the potential of natural E. huxleyi populations to cope with global climate change.

Planktonic foraminifera in the trans-tropical Pacific Ocean

Cores from four Ocean Drilling Program (ODP) sites were examined for planktonic foraminifers. One sample per core (from core-catchers in Holes 806B and 807B and from Section 4 in Holes 847B and 852B) was examined through the interval representing the last 5.8 m.y. Sites 806 (0°19.1'N; 159°21.7'E) and 847 (0o12.1'N; 95°19.2'W) are beneath the equatorial divergence zone. Sites 807 (3°36.4'N; 156°37.5'E) and 852 (5°19.6'N; 110°4.6'W) are located north of the equator in the convergence zone created by the interaction of the westward-flowing South Equatorial Current (SEC) and the eastward-flowing North Equatorial Countercurrent (NECC). Specimens were identified to species and then grouped according to depth habitat and trophic level. Species richness and diversity were also calculated. Tropical neogloboquadrinids have been more abundant in the eastern than in the western equatorial Pacific Ocean throughout the last 5.8 m.y. During the mid-Pliocene (3.8-3.2 Ma), their abundance increased at all sites, while during the Pleistocene (after ~ 1.6 Ma), they expanded in the east and declined in the west. This suggests an increase in surface-water productivity across the Pacific Ocean during the closing of the Central American seaway and an exacerbation of the productivity asymmetry between the eastern and western equatorial regions during the Pleistocene. This faunal evidence agrees with eolian grain-size data (Hovan, 1995) and diatom flux data (Iwai, this volume), which suggest increases in tradewind strength in the eastern equatorial Pacific that centered around 3.5 and 0.5 Ma. The present longitudinal zonation of thermocline dwelling species, a response to the piling of warm surface water in the western equatorial region of the Pacific, seems to have developed after 2.4 Ma, not directly after the closing of the Panama seaway (3.2 Ma). Apparently, after 2.4 Ma, the piling warm water in the west overwhelmed the upwelling of nutrients into the photic zone in that region, creating the Oceanographic asymmetry that exists in the modern tropical Pacific and is reflected in the microfossil record. In the upper Miocene and lower Pliocene sediments, the ratio of thermocline-dwelling species to mixed-layer dwellers is 60%:40%. During the mid-Pliocene, the western sites became 40% thermocline and 60% mixed-layer dwellers. Subsequent to -2.4 Ma, the asymmetry increased to 20%: 80% in the west and the reverse in the east. This documents the gradual thickening of the warm-water layer piled up in the western tropical Pacific over the last 5.8 m.y. and reveals two "steps" in the biotic trend that can be associated with specific events in the physical environment.

Initial biomass and biomass change of tundra and forest plants in three herbivore treatments in Norway and Sweden

Recent Pan-Arctic shrub expansion has been interpreted as a response to a warmer climate. However, herbivores can also influence the abundance of shrubs in arctic ecosystems. We addressed these alternative explanations by following the changes in plant community composition during the last 10 years in permanent plots inside and outside exclosures with different mesh sizes that exclude either only reindeer or all mammalian herbivores including voles and lemmings. The exclosures were replicated at three forest and tundra sites at four different locations along a climatic gradient (oceanic to continental) in northern Fennoscandia. Since the last 10 years have been exceptionally warm, we could study how warming has influenced the vegetation in different grazing treatments. Our results show that the abundance of the dominant shrub, Betula nana, has increased during the last decade, but that the increase was more pronounced when herbivores were excluded. Reindeer have the largest effect on shrubs in tundra, while voles and lemmings have a larger effect in the forest. The positive relationship between annual mean temperature and shrub growth in the absence of herbivores and the lack of relationships in grazed controls is another indication that shrub abundance is controlled by an interaction between herbivores and climate. In addition to their effects on taller shrubs (> 0.3 m), reindeer reduced the abundance of lichens, whereas microtine rodents reduced the abundance of dwarf shrubs (< 0.3 m) and mosses. In contrast to short-term responses, competitive interactions between dwarf shrubs and lichens were evident in the long term. These results show that herbivores have to be considered in order to understand how a changing climate will influence tundra ecosystems.

Gasterosteus aculeatus egg traits and fish lengths vs. experiment setup

1) Our study addresses the role of non-genetic and genetic inheritance in shaping the adaptive potential of populations under a warming ocean scenario. We used a combined experimental approach (transgenerational plasticity and quantitative genetics) to partition the relative contribution of maternal vs. paternal (additive genetic) effects to offspring body size (a key component of fitness), and investigated a potential physiological mechanism (mitochondrial respiration capacities) underlying whole organism growth/size responses. 2) In very early stages of growth (up to 30 days), offspring body size of marine sticklebacks benefited from maternal transgenerational plasticity (TGP): offspring of mothers acclimated to17°C were larger when reared at 17°C, and offspring of mothers acclimated to 21°C were larger when reared at 21°C. The benefits of maternal TGP on body size were stronger and persisted longer (up to 60 days) for offspring reared in the warmer (21°C) environment, suggesting that maternal effects will be highly relevant for climate change scenarios in this system. 3) Mitochondrial respiration capacities measured on mature offspring (F1 adults) matched the pattern of TGP for juvenile body size, providing an intuitive mechanistic basis for the maternal acclimation persisting into adulthood. Size differences between temperatures seen at early growth stages remained in the F1 adults, linking offspring body size to maternal inheritance of mitochondria. 4) Lower maternal variance components in the warmer environment were mostly driven by mothers acclimated to ambient (colder) conditions, further supporting our tenet that maternal effects were stronger at elevated temperature. Importantly, all parent-offspring temperature combination groups showed genotype x environment (GxE) interactions, suggesting that reaction norms have the potential to evolve. 5) To summarise, transgenerational plasticity and genotype x environment interactions work in concert to mediate impacts of ocean warming on metabolic capacity and early growth of marine sticklebacks. TGP can buffer short-term detrimental effects of climate warming and may buy time for genetic adaptation to catch up, therefore markedly contributing to the evolutionary potential and persistence of populations under climate change.

Sedimentology of a profile from Etoliko Lagoon, Western Greece

A key feature of Greece is the large amount of historical and archaeological records. The sedimentary record of the Etoliko Lagoon, Aetolia, Western Greece, offers an ideal opportunity to study human-environment interaction and to disentangle natural and anthropogenic imprints in the sedimentary record. By applying an interdisciplinary approach of combining geoscientific methods (XRF, LOI, grain size analysis) with archaeological and historical records, the 8.8 m long sedimentary sequence ETO1C reveals the palaeoenvironmental history of the lagoon and its catchment since 11,670 cal BP. With a thorough chronology based on 14C age-depth-modelling including varve counting, different evolutionary stages were put in a chronological context. These stages include a lake period (11,670-8310 cal BP) followed by a period of sporadic saltwater intrusion (8310-1350 cal BP) as a result of continuing transgression. Phases of limnic predominance associated with freshwater inflow of episodically activated distributaries (around 5230 cal BP) still occurred. By 1350 cal BP, ongoing sea level rise had connected the lagoons of Etoliko and Messolonghi and freshwater influence had ceased. With the onset of settlement activity in the Late Helladic (1700-1100 cal BC) humans took advantage of the prevailing environmental landscape. A sudden increase in coarse sedimentation correlates with the history of human occupation with its peak of prosperity from the Late Helladic until the end of the Hellenistic Period (30 cal BC).

Gene sequences of the lichen Cetraria aculeata

Lichens are symbioses between fungi (mycobionts) and photoautotrophic green algae or cyanobacteria (photobionts). Many lichens occupy large distributional ranges covering several climatic zones. So far, little is known about the large-scale phylogeography of lichen photobionts and their role in shaping the distributional ranges of lichens. We studied south polar, temperate and north polar populations of the widely distributed fruticose lichen Cetraria aculeata. Based on the DNA sequences from three loci for each symbiont, we compared the genetic structure of mycobionts and photobionts. Phylogenetic reconstructions and Bayesian clustering methods divided the mycobiont and photobiont data sets into three groups. An AMOVA shows that the genetic variance of the photobiont is best explained by differentiation between temperate and polar regions and that of the mycobiont by an interaction of climatic and geographical factors. By partialling out the relative contribution of climate, geography and codispersal, we found that the most relevant factors shaping the genetic structure of the photobiont are climate and a history of codispersal. Mycobionts in the temperate region are consistently associated with a specific photobiont lineage. We therefore conclude that a photobiont switch in the past enabled C. aculeata to colonize temperate as well as polar habitats. Rare photobiont switches may increase the geographical range and ecological niche of lichen mycobionts by associating them with locally adapted photobionts in climatically different regions and, together with isolation by distance, may lead to genetic isolation between populations and thus drive the evolution of lichens.

In-situ Iron isotope measurements and mineral compositions from ODP Hole 206-1256D and IODP Hole 335-U1256D

In-situ Fe isotope measurements have been carried out to estimate the impact of the hydrothermal metamorphic overprint on the Fe isotopic composition of Fe-Ti-oxides and Fe-sulfides of the different lithologies of the drilled rocks from IODP Hole 1256D (eastern equatorial Pacific; 15 Ma crust formed at the East Pacific Rise). Most igneous rocks normally have a very restricted range in their 56Fe/54Fe ratio. In contrast, Fe isotope compositions of hot fluids (> 300 °C) from mid-ocean-ridge spreading centers define a narrow range that is shifted to lower delta 56Fe values by 0.2 per mil - 0.5 per mil as compared to igneous rocks. Therefore, it is expected that mineral phases that contain large amounts of Fe are especially affected by the interaction with a fluid that fractionates Fe isotopes during exsolution/precipitation of those minerals. We have used a femtosecond UV-Laser ablation system to determine mineral 56Fe/54Fe ratios of selected samples with a precision of < 0.1 per mil (2 sigma level) at micrometer-scale. We have found significant variations of the delta 56Fe (IRMM-014) values in the minerals between different samples as well as within samples and mineral grains. The overall observed scale of delta 56Fe (magnetite) in 1256D rocks ranges from - 0.12 to + 0.64 per mil, and of delta 56Fe (ilmenite) from - 0.77 to + 0.01 per mil. Pyrite in the lowermost sheeted dike section is clearly distinguishable from the other investigated lithological units, having positive delta 56Fe values between + 0.29 and + 0.56 per mil, whereas pyrite in the other samples has generally negative delta 56Fe values from - 1.10 to - 0.59 permil. One key observation is that the temperature dependent inter-mineral fractionations of Fe isotopes between magnetite and ilmenite are systematically shifted towards higher values when compared to theoretically expected values, while synthesized, well equilibrated magnetite-ilmenite pairs are compatible with the theoretical predictions. Theoretical considerations including beta-factors of different aqueous Fe-chlorides and Rayleigh-type fractionations in the presence of a hydrous, chlorine-bearing fluid can explain this observation. The disagreement between observed and theoretical equilibrium fractionation, the fact that magnetite, in contrast to ilmenite shows a slight downhole trend in the delta 56Fe values, and the observation of small scale heterogeneities within single mineral grains imply that a general re-equilibration of the magnetite-ilmenite pairs is overprinted by kinetic fractionation effects, caused by the interaction of magnetite/ilmenite with hydrothermal fluids penetrating the upper oceanic crust during cooling, or incomplete re-equilibration at low temperatures. Furthermore, the observation of significant small-scale variations in the 56Fe/54Fe ratios of single minerals in this study highlights the importance of high spatial-resolution-analyses of stable isotope ratios for further investigations.

Geochemistry and paleontology of sediment core GeoB3359-3

A geochemical and paleontological reconstruction of paleoproductivity, upwelling intensity and sea surface temperature (SST) off central Chile at 35°S (GeoB3359-3) reveals marked changes from the Last Glacial Maximum (LGM) through the Early Holocene. Surface-water productivity was determined by the interaction between the atmospheric (the Southern Westerlies) and oceanographic (the Antarctic Circumpolar Current, ACC) systems from the LGM through early Termination I (TI). The northward shift of the climate zones during the LGM brought the ACC, as the main macronutrient source, closer to the GeoB3359-3, SST lowered, and surface water productivity and accumulation rates of biogenic components enhanced. With the poleward return of the Southern Westerlies and the ACC, the subtropical high-pressure system became the dominant atmospheric component southward till 35°S during the late TI and Early Holocene and caused surface water productivity to increase through enhanced upwelling.

Geochemistry at DSDP Leg 70 Holes

A total of 32 holes at five sites near 1°N, 86°W drilled on Deep Sea Drilling Project (DSDP) Leg 70 (November- December 1979) provide unique data on the origin of the hydrothermal mounds on the southern flank of the Galapagos Spreading Center. Hydrothermal sediments, primarily Mn-oxide and nontronite, are restricted to the immediate vicinity of the mounds (< 100 m) and are probably formed by the interaction of upward-percolating hydrothermal solutions with seawater and pelagic sediments above locally permeable zones of ocean crust. Mounds as high as 25 meters form in less than a few hundred thousand years, and geothermal and geochemical gradients indicate that they are actively forming today. The lack of alteration of upper basement rocks directly below the mounds and throughout the Galapagos region indicates that the source of the hydrothermal solutions is deeper in the crust.

Plant distribution and climate data across western North America

Using an extensive network of occurrence records for 293 plant species collected over the past 40 years across a climatically diverse geographic section of western North America, we find that plant species distributions were just as likely to shift upwards (i.e., towards higher elevations) as downward (i.e., towards lower elevations) - despite consistent warming across the study area. Although there was no clear directional response to climate warming across the entire study area, there was significant region-to region- variation in responses (i.e. from as many as 73% to as few as32% of species shifting upward or downward). To understand the factors that might be controlling region-specific distributional shifts, we explored the relationship between the direction of change in distribution limits and the nature of recent climate change. We found that the direction of distribution limit shifts was explained by an interaction between the rate of change in local summer temperatures and seasonal precipitation. Specifically, species shifted upward at their upper elevational limit when snowfall declined at slower rates and minimum temperatures increased. By contrast, species shifted upwards at their lower elevation limit when maximum temperatures increased or both temperature and precipitation decreased. Our results suggest that future species' elevational distribution shifts will be complex, depending on the interaction between seasonal temperature and precipitation change.

Granulometry, biogenic silica content, magnetic susceptibility and bulk densities of ODP Site 188-1165

The Prydz Bay area is a key region for studying and understanding the history of the eastern Antarctic Continental Ice Sheet (O'Brien, Cooper, Richter, et al., 2001, doi:10.2973/odp.proc.ir.188.2001). Ocean Drilling Program (ODP) Site 1165 is situated in a water depth of 3357 m on the continental rise offshore from Prydz Bay and lies in front of the outlet for the Lambert Glacier-Amery Ice Shelf system that today drains 22% of East Antarctica. The site was drilled into mixed pelagic and hemipelagic sediments from the southwestern side of the Wild Drift. The drift is an elongate sediment body formed by the interaction of sediment supplied from continental shelf and slope with westward-flowing bottom currents. The sedimentary sequence is characterized by alternations between a generally gray to dark gray facies and a green to greenish gray facies. The greenish facies are structureless diatom-bearing clays with common bioturbation and larger amounts (>15%-20%) of biogenic silica, dispersed clasts, and lonestones than the dark gray facies, which are mostly less bioturbated clay with some silt laminations (Shipboard Scientific Party, 2001, doi:10.2973/odp.proc.ir.188.103.2001). High-quality advanced piston corer and extended core barrel cores containing nearly complete sections of middle Miocene to early Pliocene age allow a detailed characterization of sedimentary cycles and can provide indications for ice advances of the Lambert Glacier system into Prydz Bay, for the extent of sea ice, and for changes in oceanic circulation. The purpose of this work is to provide a data set of coarse-fraction mass percentage (>63, >125, and >250 µm) and biogenic silica content measured on sediments of late Miocene to early Pliocene age drilled at Site 1165. Additionally, high-resolution records of magnetic susceptibility (MS) and gamma ray attenuation (GRA) bulk density are presented. These shipboard data sets were edited postcruise. Furthermore, I provide a high-resolution dry bulk density record that is derived from GRA bulk density and can be used for the calculation of mass accumulation rates. These sedimentological and physical parameters will be used in future work to understand the depositional pattern of alternating biogenic and terrigenous sediments that was observed at Site 1165 (Shipboard Scientific Party, 2001, doi:10.2973/odp.proc.ir.188.103.2001).

Mean sortable silt values from Holocene and glacial Scotia Sea sediments

The Antarctic Circumpolar Current is key to the mixing and ventilation of the world's oceans. This current flows from west to east between about 45° and 70° S connecting the Atlantic, Pacific and Indian oceans, and is driven by westerly winds and buoyancy forcing. High levels of productivity in the current regulate atmospheric CO2 concentrations. Reconstructions of the current during the last glacial period suggest that flow speeds were faster or similar to present, and it is uncertain whether the strength and position of the westerly winds changed. Here we reconstruct Antarctic Circumpolar Current bottom speeds through the constricting Drake Passage and Scotia Sea during the Last Glacial Maximum and Holocene based on the mean grain size of sortable silt from a suite of sediment cores. We find essentially no change in bottom flow speeds through the region, and, given that the momentum imparted by winds, and modulated by sea-ice cover, is balanced by the interaction of these flows with the seabed, this argues against substantial changes in wind stress. However, glacial flow speeds in the sea-ice zone south of 56° S were significantly slower than present, whereas flow in the north was faster, but not significantly so. We suggest that slower flow over the rough topography south of 56° S may have reduced diapycnal mixing in this region during the last glacial period, possibly reducing the diapycnal contribution to the Southern Ocean overturning circulation.

Compressional and shear wave velocities and elastic constants at DSDP Leg 59 Holes

Prior to the Deep Sea Drilling Project the composition of the oceanic crust could only be inferred from seismic-refraction and gravity data and the recovery of a wide variety of dredged rocks. Through the success of the Deep Sea Drilling Project, it is now clear that the top of oceanic Layer 2 usually consists of basalt. Several laboratory studies (e.g., Fox et al., 1972; Christensen and Shaw, 1970; Hyndman and Drury, 1976) have demonstrated that the seismic velocities of oceanic basalt are similar to velocities reported from refraction studies of Layer 2 and that the variability in Layer 2 velocities has many causes, the most important being fracturing and sea-floor alteration produced by the interaction of basalt and sea water (Christensen and Salisbury, 1973). To date, most reported measurements of velocities in oceanic basalts are from samples obtained from the main ocean basins. With the exception of an earlier study of velocities and related elastic properties of a suite of rocks from DSDP Sites 292, 293, 294, and 296 located in the Philippine Sea (Christensen et al., 1975; Fountain et al., 1975), elastic properties have not been determined for oceanic rocks from marginal basins. In this chapter compressional- and shear-wave velocities and elastic constants are reported at elevated confining pressures for basalt and volcanic breccias from Holes 447A, 448, and 448A.

Petrology, mineralogy, and chemistry of basaltic rocks at DSDP Leg 81 Holes

We detail the petrography and mineralogy of 145 basaltic rocks from the top, middle, and base of flow units identified on shipboard along with associated pyroclastic samples. Our account includes representative electron microprobe analyses of primary and secondary minerals; 28 whole-rock major-oxide analyses; 135 whole-rock analyses each for 21 trace elements; 7 whole-rock rare-earth analyses; and 77 whole-rock X-ray-diffraction analyses. These data show generally similar petrography, mineralogy, and chemistry for the basalts from all four sites; they are typically subalkaline and consanguineous with limited evolution along the tholeiite trend. Limited fractionation is indicated by immobile trace elements; some xenocrystic incorporation from more basic material also occurred. Secondary alteration products indicate early subaerial weathering followed by prolonged interaction with seawater, most likely below 150°C at Holes 552, 553A, and 554A. At Hole 555, greenschist alteration affected the deepest rocks (olivine-dolerite) penetrated, at 250-300°C.