World distribution of land cover changes, model in NetCDF format

The role of Pre- and Protohistoric anthropogenic land cover changes needs to be quantified i) to establish a baseline for comparison with current human impact on the environment and ii) to separate it from naturally occurring changes in our environment. Results are presented from the simple, adaptation-driven, spatially explicit Global Land Use and technological Evolution Simulator (GLUES) for pre-Bronze age demographic, technological and economic change. Using scaling parameters from the History Database of the Global Environment as well as GLUES-simulated population density and subsistence style, the land requirement for growing crops is estimated. The intrusion of cropland into potentially forested areas is translated into carbon loss due to deforestation with the dynamic global vegetation model VECODE. The land demand in important Prehistoric growth areas - converted from mostly forested areas - led to large-scale regional (country size) deforestation of up to 11% of the potential forest. In total, 29 Gt carbon were lost from global forests between 10 000 BC and 2000 BC and were replaced by crops; this value is consistent with other estimates of Prehistoric deforestation. The generation of realistic (agri-)cultural development trajectories at a regional resolution is a major strength of GLUES. Most of the pre-Bronze age deforestation is simulated in a broad farming belt from Central Europe via India to China. Regional carbon loss is, e.g., 5 Gt in Europe and the Mediterranean, 6 Gt on the Indian subcontinent, 18 Gt in East and Southeast Asia, or 2.3 Gt in subsaharan Africa.

Changes in coral reef communities across a natural gradient in seawater pH

Ocean acidification threatens the survival of coral reef ecosystems worldwide. The negative effects of ocean acidification observed in many laboratory experiments have been seen in studies of naturally low-pH reefs, with little evidence to date for adaptation. Recently, we reported initial data suggesting that low-pH coral communities of the Palau Rock Islands appear healthy despite the extreme conditions in which they live. Here, we build on that observation with a comprehensive statistical analysis of benthic communities across Palau's natural acidification gradient. Our analysis revealed a shift in coral community composition but no impact of acidification on coral richness, coralline algae abundance, macroalgae cover, coral calcification, or skeletal density. However, coral bioerosion increased 11-fold as pH decreased from the barrier reefs to the Rock Island bays. Indeed, a comparison of the naturally low-pH coral reef systems studied so far revealed increased bioerosion to be the only consistent feature among them, as responses varied across other indices of ecosystem health. Our results imply that whereas community responses may vary, escalation of coral reef bioerosion and acceleration of a shift from net accreting to net eroding reef structures will likely be a global signature of ocean acidification.

The effect of elevated CO2 and increased temperature on in vitro fertilization success and initial embryonic development of single male:female crosses of broad-cast spawning corals at mid- and high-latitude locations

The impact of global climate change on coral reefs is expected to be most profound at the sea surface, where fertilization and embryonic development of broadcast-spawning corals takes place. We examined the effect of increased temperature and elevated CO2 levels on the in vitro fertilization success and initial embryonic development of broadcast-spawning corals using a single male:female cross of three different species from mid- and high-latitude locations: Lyudao, Taiwan (22° N) and Kochi, Japan (32° N). Eggs were fertilized under ambient conditions (27 °C and 500 µatm CO2) and under conditions predicted for 2100 (IPCC worst case scenario, 31 °C and 1000 µatm CO2). Fertilization success, abnormal development and early developmental success were determined for each sample. Increased temperature had a more profound influence than elevated CO2. In most cases, near-future warming caused a significant drop in early developmental success as a result of decreased fertilization success and/or increased abnormal development. The embryonic development of the male:female cross of A. hyacinthus from the high-latitude location was more sensitive to the increased temperature (+4 °C) than the male:female cross of A. hyacinthus from the mid-latitude location. The response to the elevated CO2 level was small and highly variable, ranging from positive to negative responses. These results suggest that global warming is a more significant and universal stressor than ocean acidification on the early embryonic development of corals from mid- and high-latitude locations.

Acid-base balance and changes in haemolymph properties of the South African rock lobsters, Jasus lalandii, a palinurid decapod, during chronic hypercapnia

Few studies exist reporting on long-term exposure of crustaceans to hypercapnia. We exposed juvenile South African rock lobsters, Jasus lalandii, to hypercapnic conditions of pH 7.3 for 28 weeks and subsequently analysed changes in the extracellular fluid (haemolymph). Results revealed, for the first time, adjustments in the haemolymph of a palinurid crustacean during chronic hypercapnic exposure: 1) acid-base balance was adjusted and sustained by increased bicarbonate and 2) quantity and oxygen binding properties of haemocyanin changed. Compared with lobsters kept under normocapnic conditions (pH 8.0), during prolonged hypercapnia, juvenile lobsters increased bicarbonate buffering of haemolymph. This is necessary to provide optimum pH conditions for oxygen binding of haemocyanin and functioning of respiration in the presence of a strong Bohr Effect. Furthermore, modification of the intrinsic structure of the haemocyanin molecule, and not the presence of molecular modulators, seems to improve oxygen affinity under conditions of elevated pCO2.

(Table 1) Biomass and total standing crop of major wet sedge tundra species at five sites at Alexandra Fiord in 1981 and 2005

The global climate is changing rapidly and Arctic regions are showing responses to recent warming. Responses of tundra ecosystems to climate change have been examined primarily through short-term experimental manipulations, with few studies of long-term ambient change. We investigated changes in above- and belowground biomass of wet sedge tundra to the warming climate of the Canadian High Arctic over the past 25 years. Aboveground standing crop was harvested from five sedge meadow sites and belowground biomass was sampled from one of the sites in the early 1980s and in 2005 using the same methods. Aboveground biomass was on average 158% greater in 2005 than in the early 1980s. The belowground biomass was also much greater in 2005: root biomass increased by 67% and rhizome biomass by 139% since the early 1980s. Dominant species from each functional group (graminoids, shrubs and forbs) showed significant increases in aboveground biomass. Responsive species included the dominant sedge species Carex aquatilis stans, C. membranacea, and Eriophorum angustifolium, as well as the dwarf shrub Salix arctica and the forb Polygonum viviparum. However, diversity measures were not different between the sample years. The greater biomass correlated strongly with increased annual and summer temperatures over the same time period, and was significantly greater than the annual variation in biomass measured in 1980-1983. Increased decomposition and mineralization rates, stimulated by warmer soils, were likely a major cause of the elevated productivity, as no differences in the mass of litter were found between sample periods. Our results are corroborated by published short-term experimental studies, conducted in other wet sedge tundra communities which link warming and fertilization with elevated decomposition, mineralization and tundra productivity. We believe that this is the first study to show responses in High Arctic wet sedge tundra to recent climate change.

Photochemical responses of the diatom Skeletonema costatum grown under elevated CO2 concentrations to short-term changes in pH

Variability in pH is a common occurrence in many aquatic environments, due to physical, chemical and biological processes. In coastal waters, lagoons, estuaries and inland waters, pH can change very rapidly (within seconds or hours) in addition to daily and seasonal changes. At the same time, progressive ocean acidification caused by anthropogenic CO2 emissions is superimposed on these spatial and temporal pH changes. Photosynthetic organisms are therefore unavoidably subject to significant pH variations at the cell surface. Whether this will affect their response to long-term ocean acidification is still unknown, nor is it known whether the short-term sensitivity to pH change is affected by the pCO2 to which the cells are acclimated. We posed the latter open question as our experimental hypothesis: Does acclimation to seawater acidification affect the response of phytoplankton to acute pH variations? The diatom Skeletonema costatum, commonly found in coastal and estuarine waters where short-term acute changes in pH frequently occur, was selected to test the hypothesis. Diatoms were grown at both 390 (pH 8.2, low CO2; LC) and 1000 (pH 7.9, high CO2; HC) µatm CO2 for at least 20 generations, and photosynthetic responses to short-term and acute changes in pH (between 8.2 and 7.6) were investigated. The effective quantum yield of LC-grown cells decreased by ca. 70% only when exposed to pH 7.6; this was not observed when exposed to pH 7.9 or 8.2. HC-grown cells did not show significant responses in any pH treatment. Non-photochemical quenching showed opposite trends. In general, our results indicate that while LC-grown cells are rather sensitive to acidification, HC-grown cells are relatively unresponsive in terms of photochemical performance.

Coral-algae metabolism and diurnal changes in the CO2-carbonate system of bulk sea water

Precise measurements were conducted in continuous flow seawater mesocosms located in full sunlight that compared metabolic response of coral, coral-macroalgae and macroalgae systems over a diurnal cycle. Irradiance controlled net photosynthesis (Pnet), which in turn drove net calcification (Gnet), and altered pH. Pnet exerted the dominant control on [CO3]2- and aragonite saturation state (Omega arag) over the diel cycle. Dark calcification rate decreased after sunset, reaching zero near midnight followed by an increasing rate that peaked at 03:00 h. Changes in Omega arag and pH lagged behind Gnet throughout the daily cycle by two or more hours. The flux rate Pnet was the primary driver of calcification. Daytime coral metabolism rapidly removes dissolved inorganic carbon (DIC) from the bulk seawater and photosynthesis provides the energy that drives Gnet while increasing the bulk water pH. These relationships result in a correlation between Gnet and Omega arag, with Omega arag as the dependent variable. High rates of H+ efflux continued for several hours following mid-day peak Gnet suggesting that corals have difficulty in shedding waste protons as described by the Proton Flux Hypothesis. DIC flux (uptake) followed Pnet and Gnet and dropped off rapidly following peak Pnet and peak Gnet indicating that corals can cope more effectively with the problem of limited DIC supply compared to the problem of eliminating H+. Over a 24 h period the plot of total alkalinity (AT) versus DIC as well as the plot of Gnet versus Omega arag revealed a circular hysteresis pattern over the diel cycle in the coral and coral-algae mesocosms, but not the macroalgae mesocosm. Presence of macroalgae did not change Gnet of the corals, but altered the relationship between Omega arag and Gnet. Predictive models of how future global changes will effect coral growth that are based on oceanic Omega arag must include the influence of future localized Pnet on Gnet and changes in rate of reef carbonate dissolution. The correlation between Omega arag and Gnet over the diel cycle is simply the response of the CO2-carbonate system to increased pH as photosynthesis shifts the equilibria and increases the [CO3]2- relative to the other DIC components of [HCO3]- and [CO2]. Therefore Omega arag closely tracked pH as an effect of changes in Pnet, which also drove changes in Gnet. Measurements of DIC flux and H+ flux are far more useful than concentrations in describing coral metabolism dynamics. Coral reefs are systems that exist in constant disequilibrium with the water column.

Resilience to temperature and pH changes in a future climate change scenario in six strains of the polar diatom Fragilariopsis cylindrus

The effects of ocean acidification and increased temperature on physiology of six strains of the polar diatom Fragilariopsis cylindrus from Greenland were investigated. Experiments were performed under manipulated pH levels (8.0, 7.7, 7.4, and 7.1) and different temperatures (1, 5, and 8 °C) to simulate changes from present to plausible future levels. Each of the 12 scenarios was run for 7 days, and a significant interaction between temperature and pH on growth was detected. By combining increased temperature and acidification, the two factors counterbalanced each other, and therefore no effect on the growth rates was found. However, the growth rates increased with elevated temperatures by 20-50% depending on the strain. In addition, a general negative effect of increasing acidification on growth was observed. At pH 7.7 and 7.4, the growth response varied considerably among strains. However, a more uniform response was detected at pH 7.1 with most of the strains exhibiting reduced growth rates by 20-37% compared to pH 8.0. It should be emphasized that a significant interaction between temperature and pH was found, meaning that the combination of the two parameters affected growth differently than when considering one at a time. Based on these results, we anticipate that the polar diatom F. cylindrus will be unaffected by changes in temperature and pH within the range expected by the end of the century. In each simulated scenario, the variation in growth rates among the strains was larger than the variation observed due to the whole range of changes in either pH or temperature. Climate change may therefore not affect the species as such, but may lead to changes in the population structure of the species, with the strains exhibiting high phenotypic plasticity, in terms of temperature and pH tolerance towards future conditions, dominating the population.

The response of Antarctic sea ice algae to changes in pH and CO2

Ocean acidification substantially alters ocean carbon chemistry and hence pH but the effects on sea ice formation and the CO2 concentration in the enclosed brine channels are unknown. Microbial communities inhabiting sea ice ecosystems currently contribute 10-50% of the annual primary production of polar seas, supporting overwintering zooplankton species, especially Antarctic krill, and seeding spring phytoplankton blooms. Ocean acidification is occurring in all surface waters but the strongest effects will be experienced in polar ecosystems with significant effects on all trophic levels. Brine algae collected from McMurdo Sound (Antarctica) sea ice was incubated in situ under various carbonate chemistry conditions. The carbon chemistry was manipulated with acid, bicarbonate and bases to produce a pCO2 and pH range from 238 to 6066 µatm and 7.19 to 8.66, respectively. Elevated pCO2 positively affected the growth rate of the brine algal community, dominated by the unique ice dinoflagellate, Polarella glacialis. Growth rates were significantly reduced when pH dropped below 7.6. However, when the pH was held constant and the pCO2 increased, growth rates of the brine algae increased by more than 20% and showed no decline at pCO2 values more than five times current ambient levels. We suggest that projected increases in seawater pCO2, associated with OA, will not adversely impact brine algal communities.

Gene expression changes in the coccolithophore Emiliania huxleyi after 500 generations of selection to ocean acidification

Coccolithophores are unicellular marine algae that produce biogenic calcite scales and substantially contribute to marine primary production and carbon export to the deep ocean. Ongoing ocean acidification particularly impairs calcifying organisms, mostly resulting in decreased growth and calcification. Recent studies revealed that the immediate physiological response in the coccolithophore Emiliania huxleyi to ocean acidification may be partially compensated by evolutionary adaptation, yet the underlying molecular mechanisms are currently unknown. Here, we report on the expression levels of 10 candidate genes putatively relevant to pH regulation, carbon transport, calcification and photosynthesis in E. huxleyi populations short-term exposed to ocean acidification conditions after acclimation (physiological response) and after 500 generations of high CO2 adaptation (adaptive response). The physiological response revealed downregulation of candidate genes, well reflecting the concomitant decrease of growth and calcification. In the adaptive response, putative pH regulation and carbon transport genes were up-regulated, matching partial restoration of growth and calcification in high CO2-adapted populations. Adaptation to ocean acidification in E. huxleyi likely involved improved cellular pH regulation, presumably indirectly affecting calcification. Adaptive evolution may thus have the potential to partially restore cellular pH regulatory capacity and thereby mitigate adverse effects of ocean acidification.

Photophysiological responses of Southern Ocean phytoplankton to changes in CO2 concentrations: Short-term versus acclimation effects

The present study examines how different pCO2 acclimations affect the CO2- and light-dependence of photophysiological processes and O2 fluxes in four Southern Ocean (SO) key phytoplankton species. We grew Chaetoceros debilis (Cleve), Pseudo-nitzschia subcurvata (Hasle), Fragilariopsis kerguelensis (O'Meara) and Phaeocystis antarctica (Karsten) under low (160 µatm) and high (1000 ?atm) pCO2. The CO2- and light-dependence of fluorescence parameters of photosystem II (PSII) were determined by means of a fluorescence induction relaxation system (FIRe). In all tested species, nonphotochemical quenching (NPQ) is the primary photoprotection strategy in response to short-term exposure to high light or low CO2 concentrations. In C. debilis and P. subcurvata, PSII connectivity (p) and functional absorption cross-sections of PSII in ambient light (sigma PSII') also contributed to photoprotection while changes in re-oxidation times of Qa acceptor (tQa) were more significant in F. kerguelensis. The latter was also the only species being responsive to high acclimation pCO2, as these cells had enhanced relative electron transport rates (rETRs) and sigma PSII' while tQa and p were reduced under short-term exposure to high irradiance. Low CO2-acclimated cells of F. kerguelensis and all pCO2 acclimations of C. debilis and P. subcurvata showed dynamic photoinhibition with increasing irradiance. To test for the role and presence of the Mehler reaction in C. debilis and P. subcurvata, the light-dependence of O2 fluxes was estimated using membrane inlet mass spectrometry (MIMS). Our results show that the Mehler reaction is absent in both species under the tested conditions. We also observed that dark respiration was strongly reduced under high pCO2 in C. debilis while it remained unaltered in P. subcurvata. Our study revealed species-specific differences in the photophysiological responses to pCO2, both on the acclimation as well as the short-term level.

Offspring sensitivity to ocean acidification changes seasonally in a coastal marine fish

Experimental assessments of species vulnerabilities to ocean acidification are rapidly increasing in number, yet the potential for short- and long-term adaptation to high CO2 by contemporary marine organisms remains poorly understood. We used a novel experimental approach that combined bi-weekly sampling of a wild, spawning fish population (Atlantic silverside Menidia menidia) with standardized offspring CO2 exposure experiments and parallel pH monitoring of a coastal ecosystem. We assessed whether offspring produced at different times of the spawning season (April to July) would be similarly susceptible to elevated (1100 µatm, pHNIST = 7.77) and high CO2 levels (2300 µatm, pHNIST = 7.47). Early in the season (April), high CO2 levels significantly (p < 0.05) reduced fish survival by 54% (2012) and 33% (2013) and reduced 1 to 10 d post-hatch growth by 17% relative to ambient conditions. However, offspring from parents collected later in the season became increasingly CO2-tolerant until, by mid-May, offspring survival was equally high at all CO2 levels. This interannually consistent plasticity coincided with the rapid annual pH decline in the species' spawning habitat (mean pH: 1 April/31 May = 8.05/7.67). It suggests that parents can condition their offspring to seasonally acidifying environments, either via changes in maternal provisioning and/or epigenetic transgenerational plasticity (TGP). TGP to increasing CO2 has been shown in the laboratory but never before in a wild population. Our novel findings of direct CO2-related survival reductions in wild fish offspring and seasonally plastic responses imply that realistic assessments of species CO2-sensitivities must control for parental environments that are seasonally variable in coastal habitats.

Detection of a variable intracellular acid-labile carbon pool in Thalassiosira weissflogii (Heterokontophyta) and Emiliania huxleyi (Haptophyta) in response to changes in the seawater carbon system

Accumulation of an intracellular pool of carbon (C(i) pool) is one strategy by which marine algae overcome the low abundance of dissolved CO2 (CO2 (aq) ) in modern seawater. To identify the environmental conditions under which algae accumulate an acid-labile C(i) pool, we applied a (14) C pulse-chase method, used originally in dinoflagellates, to two new classes of algae, coccolithophorids and diatoms. This method measures the carbon accumulation inside the cells without altering the medium carbon chemistry or culture cell density. We found that the diatom Thalassiosira weissflogii [(Grunow) G. Fryxell & Hasle] and a calcifying strain of the coccolithophorid Emiliania huxleyi [(Lohmann) W. W. Hay & H. P. Mohler] develop significant acid-labile C(i) pools. C(i) pools are measureable in cells cultured in media with 2-30 µmol/l CO2 (aq), corresponding to a medium pH of 8.6-7.9. The absolute C(i) pool was greater for the larger celled diatoms. For both algal classes, the C(i) pool became a negligible contributor to photosynthesis once CO2 (aq) exceeded 30 µmol/l. Combining the (14) C pulse-chase method and (14) C disequilibrium method enabled us to assess whether E. huxleyi and T. weissflogii exhibited thresholds for foregoing accumulation of DIC or reduced the reliance on bicarbonate uptake with increasing CO2 (aq) . We showed that the C(i) pool decreases with higher CO2 :HCO3 (-) uptake rates.

Biomineralization changes with food supply confer juvenile scallops (Argopecten purpuratus) resistance to ocean acidification

Future ocean acidification (OA) will affect physiological traits of marine species, with calcifying species being particularly vulnerable. As OA entails high energy demands, particularly during the rapid juvenile growth phase, food supply may play a key role in the response of marine organisms to OA. We experimentally evaluated the role of food supply in modulating physiological responses and biomineralization processes in juveniles of the Chilean scallop, Argopecten purpuratus, that were exposed to control (pH 8.0) and low pH (pH 7.6) conditions using three food supply treatments (high, intermediate, and low). We found that pH and food levels had additive effects on the physiological response of the juvenile scallops. Metabolic rates, shell growth, net calcification, and ingestion rates increased significantly at low pH conditions, independent of food. These physiological responses increased significantly in organisms exposed to intermediate and high levels of food supply. Hence, food supply seems to play a major role modulating organismal response by providing the energetic means to bolster the physiological response of OA stress. On the contrary, the relative expression of chitin synthase, a functional molecule for biomineralization, increased significantly in scallops exposed to low food supply and low pH, which resulted in a thicker periostracum enriched with chitin polysaccharides. Under reduced food and low pH conditions, the adaptive organismal response was to trade-off growth for the expression of biomineralization molecules and altering of the organic composition of shell periostracum, suggesting that the future performance of these calcifiers will depend on the trajectories of both OA and food supply. Thus, incorporating a suite of traits and multiple stressors in future studies of the adaptive organismal response may provide key insights on OA impacts on marine calcifiers.