Pollen records and lithology of profiles from Lobsigensee, Switzerland

Lobsigensee is a small kettle hole lake 15 km north-west of Bern on the Swiss Plateau, at an altitude of 514 m asl. Its surface is 2ha today, its maximum depth 2.7 m; it has no inlet and the overflow functions mainly during snow melting. The area was covered by Rhone ice during the Last Glaciation (map in Fig.2). Local geology, climate and vegetation are summarized in Figure 3A-C, the history of settlement in Figures 5-7. In order to reconstruct the vegetational and environmental history of the lake and its surroundings pollen analysis and other bio- and isotope stratigraphies were applied to twelve profiles cored across the basin with modified Livingstone corers (Fig.3 D). (1) The standard diagram: The central core LQ-90 is described as the standard pollen diagram (Chapter 3) with 10 local pollen assemblage zones of the Late-Glacial (local PAZ Ll to Ll0, from about 16'000(7) to 10'000 years BP) and 20 PAZ of the Holocene (local PAZ L11 to L30), see Figs. 8-10 and 20-24. Local PAZ L 1 to L3 are in the Late-Glacial clay and record the vegetational development after the ice retreat: L1 shows very low pollen concentration and high Pinus percentages due to long-distance transport and reworking; the latter mechanism is corroborated by the findings of thermophilous and pre-Quaternary taxa. Local PAZ L2 has a high di versi ty of non-arboreal pollen (NAP) and reflects the Late-Glacial steppe rich in heliophilous species. Local PAZ L3 is similar but additionally rich in Betula nana and Sal1x, thus reflecting a "shrub tundra". The PAZ L1 to L3 belong to the Oldest Dryas biozone. Local PAZ L4 to L 10 are found in the gyttja of the profundal or in the lake marl of the littoral and record the Late-Glacial forests. L4 is the shrub phase of reforestation with very high Junlperus and rapidly increasing Betula percentages. L5 is the PAZ with a first, L7 with a second dominance of tree-birches, separated by L6 showing a depression in the Betula curve. L4 to L7 can be assigned to the Balling biozone. Possible correlation of the Betula depression to the Older Dryas biozone is discussed. In local PAZ L8 Plnus immigrates and expands. L9 shows a facies difference in that Plnus dominates over Betula in littoral but not in profundal spectra. L8 and L9 belong to the Allerod biozone. In its youngest part the volcanic ash from Laach/Eifel is regularly found (11,000 BP). The local PAZ Ll0 corresponds to the Younger Dryas blozone. The merely slight increase of the NAP indicates that the pine forests of the lowland were not strongly affected by a cooler climate. In order to evaluate the significance of the littoral accumulation of coniferous pollen the littoral profile LQ-150 is compared to the profundal. Radiocarbon stratigraphies derived from different materials are presented in Figures 13 and 14 and in Tables 2 and 3. The hard-water errors in the gyttja samples and the carbonate samples are similar. The samples of terrestrial plant macrofossils are not affected by hard-water errors. Two plateaux of constant age appear in the age-depth relationship; their consequence for biostratigraphy as well as pollen concentration and influx diagrams are discussed. Radiocarbon ages of the Late-Glacial pollen zones are shown in Table 10. The Holocene vegetational history is recorded in the local PAZ L 11 to L30. After a Preboreal (PAZ L11) dominated by pine and birch the expansions of Corylus, Ulmus and Quercus are very rapid. Among these taxa Corylus dominates dur ing the Boreal (PAZ L 12 and L 1 3), whereas the components of the mixed oak forest dominate in the Older Atlantic (PAZ L14 to L16). In the Younger Atlantic (PAZ L 17 to L 19) Fagus and Alnus play an increasing, the mixed oak forest a decreasing role. During the period of local PAZ L19 Neolithic settlers lived on the shore of Lobsigensee. During the Subboreal (PAZ L20 and L21) and the Older Subatlantic (L22 to L25) strong fluctuations of Fagus and often antagonistic peaks of NAP, Alnus, Betula and Corylus can be interpreted as signs of human impact on vegetation. L23 is characterized not only by high values of NAP (especially apophytes and anthropochorous species) but also by the appearance of Juglans, Castanea and Secale which point to the Roman colonization of the area. For a certain period during the Younger Subatlantic (PAZ L26 to L30) the lake was used for retting hemp (Cannabis). Later the dominance of Quercus pollen indicates the importance of wood pastures. The youngest sediments reflect the wide-spread agricultural grass lands and the plantation of Pinus and Picea. Radiocarbon dates for the Holocene are given in Figure 23 and Table 4, the extrapolated ages of the Holocene pollen zones in Table 15. (2) The cross sections: Figures 25 and 26 give a summary of the litho- and palynostratigraphy of the two cross sections. Based on 11 Late-Glacial and 9 Holocene pollen diagrams (in addition to the standard ones), the consistency of the criteria for the definition of the pollen zones is examined in Tables 7 and 8 for the Late-Glacial and in Tables 11 to 14 for the Holocene. Sediment thicknesses across the basin for each pollen zone are presented in these tables as well as in Figures 43 to 45 for the Late-Glacial and in Figures 59 to 65 for the Holocene. Sediment focusing can explain differences between the gyttja cores of the profundal. Focusing is more than compensated for through "stretching" by carbonate precipitation on the littoral terrace. Pollen influx to the cross section are discussed (Chapters 4.1.5. and 4.2.3.). (3) The regional pollen zones: Based on some selected sites between Lake Geneva and Lake Constance regional pollen zones are proposed (Table 16, 17 and 19). (4) Paleoecology: Climatic change in the Late-Glacial can be inferred from Coleoptera, Trichoptera, Chironomidae and d18O of carbonates: a distinct warming is recorded around 12' 600 BP and around 10' 000 BP. The Younger Dryas biozone (10'700-10'000 BP) was the only cooling found in the Late-Glacial. The Betula depression often correlated wi th the Older Dryas biozone was possibl not colder but dryer than the previous period. During the Holocene the lowland site is not very sensitive to the minor climatic changes. Table 22 summarizes climatic and trophic changes before 8'000 BP as deduced from various biostratigraphies studied by a number of authors. Ostracods, Chironomids and fossil pigments indicate that anoxic conditions prevailed during the BoIling (possibly meromixis). Changes in the lake level are illustrated in Figure 74. A first lake-level lowering occurred in the early Holocene (10'000 to 9'000 BP), a second during the Atlantic (about 6'800 to 5'200 BP). The first "shrinking" of the lake volume resulted in a eutrophication recorded by laminations in the profundal and by pigments of Cyanophyceae. The second fall in water level corresponds to an increase of Nymphaeaceae. Human impact can be inferred in three ways: eutrophication of the lake (since the Neolithic), changes of terrestrial vegetation by deforestations (cyclicity of Fagus, see Figures 78 to 80), and enhanced erosion (increasing sedimentation rates by inwashed clay, particularly since the Roman Colonization, see Figures 49 and 81). Summary: This paper was planned as the final report on Lobsigensee. However, a number of issues are not answered but can only be asked more precisely, for example: (1) For the two periods with the highest rates of change, Le. the Bolling and the Preboreal biozones, pollen influx may reflect vegetation dynamics. Detailed investigations of these periods in annually laminated sediments are planned. (2) Biostratigraphies other than palynostratigraphy are needed to estimate the degree of linkage or independence in the development of terrestrial and lacustrine ecosystems. Often our sampling intervals were not identical, thus influencing our temporal resolution. (3) 6180- and 14C-stratigraPhies with high resolution will elucidate the leads and lags of these dynamic periods. Plateaux of constant age in the age-depth relationship have a strong bearing on both biological and geophysical understanding of Late-Glacial and early Holocene developments. (4) Numerical methods applied to the pollen diagrams of the cross section will help to quantify the significance of similari ties and dissimilarities across a single basin (with Prof. Birks). (5) Numerical methods applied to different sites on the Swiss Plateau and on the transect across the Alps will be helpful in evaluating the influence of different environmental factors (with Prof. Birks). (6) A new map 1: 1000 with 50cm-contour lines prov ided by Prof. Zurbuchen will be combined with a grid of cores sampling the transition from lake marl to peat enabling us to calculate paleo-volumes of the lake. This is interesting for the two "shrinking periods" (in Fig. 74A numbers 2-6 and 7-10), both accompanied by eutrophication. The pal eo-volume during the Neoli thic set tlement of the Cortaillod culture linked wi th an est l.mate of trophic change derived from diatoms (Prof. Smol in prep.) could possibly give an indication of the size of the human population of this period. (7) For the period with the antagonism between Fagus peaks and ABC-peaks close collaboration between palynologists, geochemists and archeologists should enable us to determine the influence of prehistoric and historic people on vegetation (collaboration with Prof. Stockli and Prof. Herzig). (8) The core LL-75 taken with a "cold letter box" will be analysed for major and trace elements by Dr. Sturm for 210pb and 137Cs by Prof.von Gunten and for pollen. We will see if our local PAZ L30 really corresponds to the surface sediment and if the small seepage lake reflects modern pollution.

Iron limitation modulates ocean acidification effects on southern ocean phytoplankton communities

The potential interactive effects of iron (Fe) limitation and Ocean Acidification in the Southern Ocean (SO) are largely unknown. Here we present results of a long-term incubation experiment investigating the combined effects of CO2 and Fe availability on natural phytoplankton assemblages from the Weddell Sea, Antarctica. Active Chl a fluorescence measurements revealed that we successfully cultured phytoplankton under both Fe-depleted and Fe-enriched conditions. Fe treatments had significant effects on photosynthetic efficiency (Fv/Fm; 0.3 for Fe-depleted and 0.5 for Fe-enriched conditions), non-photochemical quenching (NPQ), and relative electron transport rates (rETR). pCO2 treatments significantly affected NPQ and rETR, but had no effect on Fv/Fm. Under Fe limitation, increased pCO2 had no influence on C fixation whereas under Fe enrichment, primary production increased with increasing pCO2 levels. These CO2-dependent changes in productivity under Fe-enriched conditions were accompanied by a pronounced taxonomic shift from weakly to heavily silicified diatoms (i.e. from Pseudo-nitzschia sp. to Fragilariopsis sp.). Under Fe-depleted conditions, this functional shift was absent and thinly silicified species dominated all pCO2 treatments (Pseudo-nitzschia sp. and Synedropsis sp. for low and high pCO2, respectively). Our results suggest that Ocean Acidification could increase primary productivity and the abundance of heavily silicified, fast sinking diatoms in Fe-enriched areas, both potentially leading to a stimulation of the biological pump. Over much of the SO, however, Fe limitation could restrict this possible CO2 fertilization effect.

Physiological responses of the marine diatom Thalassiosira pseudonana to increased pCO2 and seawater acidity

We studied the effects of elevated CO2 concentration and seawater acidity on inorganic carbon acquisition, photoinhibition and photoprotection as well as growth and respiration in the marine diatom Thalassiosira pseudonana. After having grown under the elevated CO2 level (1000 µatm, pH 7.83) at sub-saturating photosynthetically active radiation (PAR, 75 µmol photons/m**2/s) for 20 generations, photosynthesis and dark respiration of the alga increased by 25% (14.69 ± 2.55 fmol C/cell/h) and by 35% (4.42 ± 0.98 fmol O2/cell/h), respectively, compared to that grown under the ambient CO2 level (390 µatm, pH 8.16), leading to insignificant effects on growth (1.09 ± 0.08 (1/d))v 1.04 ± 0.07 (1/d)). The photosynthetic affinity for CO2 was lowered in the high-CO2 grown cells, reflecting a down-regulation of the CO2 concentrating mechanism (CCM). When exposed to an excessively high level of PAR, photochemical and non-photochemical quenching responded similarly in the low- and high-CO2 grown cells, reflecting that photoinhibition was not influenced by the enriched level of CO2. In T. pseudonana, it appeared that the energy saved due to the down-regulated CCM did not contribute to any additional light stress as previously found in another diatom Phaeodactylum tricornutum, indicating differential physiological responses to ocean acidification between these two diatom species.

Seawater carbonate chemistry and photosynthesis of a coccolithophorid in a laboratory experiment

Mixing of seawater subjects phytoplankton to fluctuations in photosynthetically active radiation (400-700 nm) and ultraviolet radiation (UVR; 280-400 nm). These irradiance fluctuations are now superimposed upon ocean acidification and thinning of the upper mixing layer through stratification, which alters mixing regimes. Therefore, we examined the photosynthetic carbon fixation and photochemical performance of a coccolithophore, Gephyrocapsa oceanica, grown under high, future (1,000 µatm) and low, current (390 µatm) CO2 levels, under regimes of fluctuating irradiances with or without UVR. Under both CO2 levels, fluctuating irradiances, as compared with constant irradiance, led to lower nonphotochemical quenching and less UVR-induced inhibition of carbon fixation and photosystem II electron transport. The cells grown under high CO2 showed a lower photosynthetic carbon fixation rate but lower nonphotochemical quenching and less ultraviolet B (280-315 nm)-induced inhibition. Ultraviolet A (315-400 nm) led to less enhancement of the photosynthetic carbon fixation in the high-CO2-grown cells under fluctuating irradiance. Our data suggest that ocean acidification and fast mixing or fluctuation of solar radiation will act synergistically to lower carbon fixation by G. oceanica, although ocean acidification may decrease ultraviolet B-related photochemical inhibition.

Properties of seawater and particulate matter from a FRRF instrument, operating in a flow-through mode on the continuous surface water sampling system during the Tara Oceans expedition 2009-2013

The Tara Oceans Expedition (2009-2013) sampled the world oceans on board a 36 m long schooner, collecting environmental data and organisms from viruses to planktonic metazoans for later analyses using modern sequencing and state-of-the-art imaging technologies. Tara Oceans Data are particularly suited to study the genetic, morphological and functional diversity of plankton. The present data set provides continuous measurements made with a FRRF instrument, operating in a flow-through mode during the 2009-2012 part of the expedition. It operates by exciting chlorophyll fluorescence using a series of short flashes of controlled energy and time intervals (Kolber et al, 1998). The fluorescence transients produced by this excitation signal were analysed in real-time to provide estimates of abundance of photosynthetic pigments, the photosynthetic yields (Fv/Fm), the functional absorption cross section (a proxy for efficiency of photosynthetic energy acquisition), the kinetics of photosynthetic electron transport between Photosystem II and Photosystem I, and the size of the PQ pool. These parameters were measured at excitation wavelength of 445 nm, 470nm, 505 nm, and 535 nm, allowing to assess the presence and the photosynthetic performance of different phytoplankton taxa based on the spectral composition of their light harvesting pigments. The FRRF-derived photosynthetic characteristics were used to calculate the initial slope, the half saturation, and the maximum level of Photosynthesis vs Irradiance relationship. FRRF data were acquired continuously, at 1-minute time intervals.

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.

Seawater carbonate chemistry and combined mechanistic effects of CO2 and light on the N2-fixing cyanobacterium Trichodesmium IMS101, 2010

The marine diazotrophic cyanobacterium Trichodesmium responds to elevated atmospheric CO2 partial pressure (pCO2) with higher N2 fixation and growth rates. To unveil the underlying mechanisms, we examined the combined influence of pCO2(150 and 900 µatm) and light (50 and 200 µmol photons m-2 s-1) on TrichodesmiumIMS101. We expand on a complementary study that demonstrated that while elevated pCO2 enhanced N2 fixation and growth, oxygen evolution and carbon fixation increased mainly as a response to high light. Here, we investigated changes in the photosynthetic fluorescence parameters of photosystem II, in ratios of the photosynthetic units (photosystem I:photosystem II), and in the pool sizes of key proteins involved in the fixation of carbon and nitrogen as well as their subsequent assimilation. We show that the combined elevation in pCO2 and light controlled the operation of the CO2-concentrating mechanism and enhanced protein activity without increasing their pool size. Moreover, elevated pCO2 and high light decreased the amounts of several key proteins (NifH, PsbA, and PsaC), while amounts of AtpB and RbcL did not significantly change. Reduced investment in protein biosynthesis, without notably changing photosynthetic fluxes, could free up energy that can be reallocated to increase N2 fixation and growth at elevated pCO2 and light. We suggest that changes in the redox state of the photosynthetic electron transportchain and posttranslational regulation of key proteins mediate the high flexibility in resources and energy allocation in Trichodesmium. This strategy should enableTrichodesmium to flourish in future surface oceans characterized by elevated pCO2, higher temperatures, and high light.

Seawater carbonate chemistry and the photophysiology of Thalassiosira pseudonana (Bacillariophyceae) and Emiliania huxleyi (Haptophyta) in a laboratory experiment

Increasing anthropogenic carbon dioxide is causing changes to ocean chemistry, which will continue in a predictable manner. Dissolution of additional atmospheric carbon dioxide leads to increased concentrations of dissolved carbon dioxide and bicarbonate and decreased pH in ocean water. The concomitant effects on phytoplankton ecophysiology, leading potentially to changes in community structure, are now a focus of concern. Therefore, we grew the coccolithophore Emiliania huxleyi (Lohmann) W. W. Hay et H. Mohler and the diatom strains Thalassiosira pseudonana (Hust.) Hasle et Heimdal CCMP 1014 and T. pseudonana CCMP 1335 under low light in turbidostat photobioreactors bubbled with air containing 390 ppmv or 750 ppmv CO2. Increased pCO2 led to increased growth rates in all three strains. In addition, protein levels of RUBISCO increased in the coastal strains of both species, showing a larger capacity for CO2 assimilation at 750 ppmv CO2. With increased pCO2, both T. pseudonana strains displayed an increased susceptibility to PSII photoinactivation and, to compensate, an augmented capacity for PSII repair. Consequently, the cost of maintaining PSII function for the diatoms increased at increased pCO2. In E. huxleyi, PSII photoinactivation and the counter-acting repair, while both intrinsically larger than in T. pseudonana, did not change between the current and high-pCO2 treatments. The content of the photosynthetic electron transport intermediary cytochrome b6/f complex increased significantly in the diatoms under elevated pCO2, suggesting changes in electron transport function.

(Table 1) Results of pore water analyses, including boron content and boron isotope ratios

Drilling a transect of holes across the Costa Rica forearc during ODP Leg 170 demonstrated the margin wedge to be of continental, non accretionary origin, which is intersected by permeable thrust faults. Pore waters from four drillholes, two of which penetrated the décollement zone and reached the underthrust lower plate sedimentary sequence of the Cocos Plate, were examined for boron contents and boron isotopic signatures. The combined results show dilution of the uppermost sedimentary cover of the forearc, with boron contents lower than half of the present-day seawater values. Pore fluid "refreshening" suggests that gas hydrate water has been mixed with the sediment interstitial water, without profoundly affecting the d11B values. Fault-related flux of a deeply generated fluid is inferred from high B concentration in the interval beneath the décollement, being released from the underthrust sequence with incipient burial. First-order fluid budget calculations over a cross-section across the Costa Rica forearc indicate that no significant fluid transfer from the lower to the upper plate is inferred from boron fluid profiles, at least within the frontal 40 km studied. Expulsed lower plate pore water, which is estimated to be 0.26-0.44 km3 per km trench, is conducted efficiently along and just beneath the décollement zone, indicating effective shear-enhanced compaction. In the upper plate forearc wedge, dewatering occurs as diffuse transport as well as channelled flow. A volume of approximately 2 km3 per km trench is expulsed due to compaction and, to a lesser extent, lateral shortening. Pore water chemistry is influenced by gas hydrate instability, so that it remains unknown whether deep processes like mineral dehydration or hydrocarbon formation may play a considerable role towards the hinterland.

Reduced calcification decreases photoprotective capability in the Coccolithophorid Emiliania huxleyi

Intracellular calcification of coccolithophores generates CO2 and consumes additional energy for acquisition of calcium and bicarbonate ions; therefore, it may correlate with photoprotective processes by influencing the energetics. To address this hypothesis, a calcifying Emiliania huxleyi strain (CS-369) was grown semi-continuously at reduced (0.1 mM, LCa) and ambient Ca2+ concentrations (10 mM, HCa) for 150 d (>200 generations). The HCa-grown cells had higher photosynthetic and calcification rates and higher contents of Chl a and carotenoids compared with the naked (bearing no coccoliths) LCa-grown cells. When exposed to stressfull levels of photosynthetically active radiation (PAR), LCa-grown cells displayed lower photochemical yield and less efficient non-photochemical quenching (NPQ). When the LCa- or HCa-grown cells were inversely shifted to their counterpart medium, LCa to HCa transfer increased photosynthetic carbon fixation (P), calcification rate (C), the C/P ratio, NPQ and pigment contents, whereas those shifted from HCa to LCa exhibited the opposite effects. Increased NPQ, carotenoids and quantum yield were clearly linked with increased or sustained calcification in E. huxleyi. The calcification must have played a role in dissipating excessive energy or as an additional drainage of electrons absorbed by the photosynthetic antennae. This phenomenon was further supported by testing two non-calcifying strains, which showed insignificant changes in photosynthetic carbon fixation and NPQ when transferred to LCa conditions