Lake-level rise in the late Pleistocene and active subaquatic volcanism since the Holocene in Lake Kivu; East African Rift

The history of Lake Kivu is strongly linked to the activity of the Virunga volcanoes. Subaerial and subaquatic volcanoes, in addition to lake-level changes, shape the subaquatic morphologic and structural features in Lake Kivu's Main Basin. Previous studies revealed that volcanic eruptions blocked the former outlet of the lake to the north in the late Pleistocene, leading to a substantial rise in the lake level and subsequently the present- day thermohaline stratification. Additional studies have speculated that volcanic and seismic activities threaten to trigger a catastrophic release of the large amount of gases dissolved in the lake. The current study presents a bathymetric mapping and seismic profiling survey that covers the volcanically active area of the Main Basin at a resolution that is unprecedented for Lake Kivu. New geomorphologic features identified on the lake floor can accurately describe related lake-floor processes for the first time. The late Pleistocene lowstand is observed at 425 m depth, and volcanic cones, tuff rings, and lava flows observed above this level indicate both subaerial and subaquatic volcanic activities during the Holocene. The geomorphologic analysis yields new implications on the geologic processes that have shaped Lake Kivu's basin, and the presence of young volcanic features can be linked to the possibility of a lake overturn.

What determines the impact of alien birds and mammals in Europe?

An often-cited reason for studying the process of invasion by alien species is that the understanding sought can be used to mitigate the impacts of the invaders. Here, we present an analysis of the correlates of local impacts of established alien bird and mammal species in Europe, using a recently described metric to quantify impact. Large-bodied, habitat generalist bird and mammal species that are widespread in their native range, have the greatest impacts in their alien European ranges, supporting our hypothesis that surrogates for the breadth and the amount of resources a species uses are good indicators of its impact. However, not all surrogates are equally suitable. Impacts are generally greater for mammal species giving birth to larger litters, but in contrast are greater for bird species laying smaller clutches. There is no effect of diet breadth on impacts in birds or mammals. On average, mammals have higher impacts than birds. However, the relationships between impact and several traits show common slopes for birds and mammals, and relationships between impact and body mass and latitude do not differ between birds and mammals. These results may help to anticipate which species would have large impacts if introduced, and so direct efforts to prevent such introductions.

Holocene climate, fire and vegetation dynamics at the treeline in the Northwestern Swiss Alps

Treelines are expected to rise to higher elevations with climate warming; the rate and extent however are still largely unknown. Here we present the first multi-proxy palaeoecological study from the treeline in the Northwestern Swiss Alps that covers the entire Holocene. We reconstructed climate, fire and vegetation dynamics at Iffigsee, an alpine lake at 2,065 m a.s.l., by using seismic sedimentary surveys, loss on ignition, visible spectrum reflectance spectroscopy, pollen, spore, macrofossil and charcoal analyses. Afforestation with Larix decidua and tree Betula (probably B. pendula) started at ~9,800 cal. b.p., more than 1,000 years later than at similar elevations in the Central and Southern Alps, indicating cooler temperatures and/or a high seasonality. Highest biomass production and forest position of ~2,100–2,300 m a.s.l. are inferred during the Holocene Thermal Maximum from 7,000 to 5,000 cal. b.p. With the onset of pastoralism and transhumance at 6,800–6,500 cal. b.p., human impact became an important factor in the vegetation dynamics at Iffigsee. This early evidence of pastoralism is documented by the presence of grazing indicators (pollen, spores), as well as a wealth of archaeological finds at the nearby mountain pass of Schnidejoch. Human and fire impact during the Neolithic and Bronze Ages led to the establishment of pastures and facilitated the expansion of Picea abies and Alnus viridis. We expect that in mountain areas with land abandonment, the treeline will react quickly to future climate warming by shifting to higher elevations, causing drastic changes in species distribution and composition as well as severe biodiversity losses.

A meta-analysis of cambium phenology and growth: linear and non-linear patterns in conifers of the northern hemisphere

Background and Aims Ongoing global warming has been implicated in shifting phenological patterns such as the timing and duration of the growing season across a wide variety of ecosystems. Linear models are routinely used to extrapolate these observed shifts in phenology into the future and to estimate changes in associated ecosystem properties such as net primary productivity. Yet, in nature, linear relationships may be special cases. Biological processes frequently follow more complex, non-linear patterns according to limiting factors that generate shifts and discontinuities, or contain thresholds beyond which responses change abruptly. This study investigates to what extent cambium phenology is associated with xylem growth and differentiation across conifer species of the northern hemisphere. Methods Xylem cell production is compared with the periods of cambial activity and cell differentiation assessed on a weekly time scale on histological sections of cambium and wood tissue collected from the stems of nine species in Canada and Europe over 1–9 years per site from 1998 to 2011. Key Results The dynamics of xylogenesis were surprisingly homogeneous among conifer species, although dispersions from the average were obviously observed. Within the range analysed, the relationships between the phenological timings were linear, with several slopes showing values close to or not statistically different from 1. The relationships between the phenological timings and cell production were distinctly non-linear, and involved an exponential pattern. Conclusions The trees adjust their phenological timings according to linear patterns. Thus, shifts of one phenological phase are associated with synchronous and comparable shifts of the successive phases. However, small increases in the duration of xylogenesis could correspond to a substantial increase in cell production. The findings suggest that the length of the growing season and the resulting amount of growth could respond differently to changes in environmental conditions.

Are ectomycorrhizal fungi alleviating or aggravating nitrogen limitation of tree growth in boreal forests?

•Symbioses between plant roots and mycorrhizal fungi are thought to enhance plant uptake of nutrients through a favourable exchange for photosynthates. Ectomycorrhizal fungi are considered to play this vital role for trees in nitrogen (N)-limited boreal forests. •We followed symbiotic carbon (C)–N exchange in a large-scale boreal pine forest experiment by tracing 13CO2 absorbed through tree photosynthesis and 15N injected into a soil layer in which ectomycorrhizal fungi dominate the microbial community. •We detected little 15N in tree canopies, but high levels in soil microbes and in mycorrhizal root tips, illustrating effective soil N immobilization, especially in late summer, when tree belowground C allocation was high. Additions of N fertilizer to the soil before labelling shifted the incorporation of 15N from soil microbes and root tips to tree foliage. •These results were tested in a model for C–N exchange between trees and mycorrhizal fungi, suggesting that ectomycorrhizal fungi transfer small fractions of absorbed N to trees under N-limited conditions, but larger fractions if more N is available. We suggest that greater allocation of C from trees to ectomycorrhizal fungi increases N retention in soil mycelium, driving boreal forests towards more severe N limitation at low N supply.

Fruit production in three masting tree species does not rely on stored carbon reserves

Fruiting is typically considered to massively burden the seasonal carbon budget of trees. The cost of reproduction has therefore been suggested as a proximate factor explaining observed mast-fruiting patterns. Here, we used a large-scale, continuous 13C labeling of mature, deciduous trees in a temperate Swiss forest to investigate to what extent fruit formation in three species with masting reproduction behavior (Carpinus betulus, Fagus sylvatica, Quercus petraea) relies on the import of stored carbon reserves. Using a free-air CO2 enrichment system, we exposed trees to 13C-depleted CO2 during 8 consecutive years. By the end of this experiment, carbon reserve pools had significantly lower δ13C values compared to control trees. δ13C analysis of new biomass during the first season after termination of the CO2 enrichment allowed us to distinguish the sources of built-in carbon (old carbon reserves vs. current assimilates). Flowers and expanding leaves carried a significant 13C label from old carbon stores. In contrast, fruits and vegetative infructescence tissues were exclusively produced from current, unlabeled photoassimilates in all three species, including F. sylvatica, which had a strong masting season. Analyses of δ13C in purified starch from xylem of fruit-bearing shoots revealed a complete turn-over of starch during the season, likely due to its usage for bud break. This study is the first to directly demonstrate that fruiting is independent from old carbon reserves in masting trees, with significant implications for mechanistic models that explain mast seeding.

PREPARATION AND CHARACTERIZATION OF COVALENTLY LINKED ENZYME-ANTIBODY CONJUGATES AND THEIR USE IN MEASURING MINUTE QUANTITIES OF PROTEIN ANTIGENS

The discovery and characterization of oncofetal proteins have led to significant advances in early cancer diagnosis and therapeutic monitoring of patients undergoing cancer chemotherapy. These tumor-associated antigens are presently measured by sensitive, specific immunoassay techniques based on the detection of minute amounts of labeled antigen or antibody incorporated into immune complexes, which must be isolated from free antigen and antibody.^ Since there are several disadvantages with using radioisotopes, the most common immunolabel, one major objective was to prepare covalently coupled enzyme-antibody conjugates and evaluate their use as a practical alternative to radiolabeled immune reagents. An improved technique for the production of enzyme-antibody conjugates was developed that involves oxidizing the carbohydrate moieties on a glycoprotein enzyme, then introducing antibody in the presence of polyethylene glycol (PEG). Covalent enzyme-antibody conjugates involving alkaline phosphatase and amyloglucosidase were produced and characterized.^ In order to increase the sensitivity of detecting the amyloglucosidase-antibody conjugate, an enzyme cycling assay was developed that measures glucose, the product of maltose cleavage by amyloglucosidase, in the picomole range. The increased sensitivity obtained by combined usage of the amyloglucosidase-antibody conjugate and enzyme cycling assay was then compared to that of conventional enzyme immunoassay (EIA).^ For immune complex isolation, polystyrene tubes and protein A-bearing Staphylococcus aureus were evaluated as solid phase matrices, upon which antibodies can be immobilized. A sandwich-type EIA, using antibody-coated S. aureus, was developed that measures human albumin (HSA) in the nanogram range. The assay, using an alkaline phosphatase-anti-HSA conjugate, was applied to the determination of HSA in human urine and evaluated extensively for its clinical applicability.^ Finally, in view of the clinical significance of alpha-fetoprotein (AFP) as an oncofetal antigen and the difficulty with its purification for use as an immunogen and assay standard, a chemical purification protocol was developed that resulted in a high yield of immunochemically pure AFP. ^

The potential of stomata analysis in conifers to estimate presence of conifer trees: examples from the Alps

To estimate whether or not a plant taxon found in the fossil record was locally present may be difficult if only pollen is analyzed. Plant macrofossils, in contrast, provide a clear indication of a taxon's local presence, although in some lake sediments or peats, macrofossils may be rare or degraded. For conifers, the stomata found on pollen slides are derived from needles and thus provide a valuable proxy for local presence and they can be identified to genus level. From previously published studies, a transect across the Alps based on 13 sites is presented. For basal samples in sandy silt above the till with high pollen values of Pinus, for example, we may distinguish pine pollen from distant sources (samples with no stomata), from reworked pollen (samples with stomata present). The first apparent local presence of most conifer genera based on stomata often but not always occurs together with the phase of rapid pollen increase (rational limit). An exception is Larix, with its annual deposition of needles and heavy poorly dispersed pollen, for it often shows the first stomata earlier, at the empirical pollen limit. The decline and potential local extinction of a conifer can sometimes be shown in the stomata record. The decline may have been caused by climatic change, competition, or human impact. In situations where conifers form the timberline, the stomata record may indicate timberline fluctuations. In the discussion of immigration or migration of taxa we advocate the use of the cautious term ``apparent local presence'' to include some uncertainties. Absence of a taxon is impossible to prove.

More efficient aboveground nitrogen use in more diverse Central European forest canopies

We hypothesized that biodiversity improves ecosystem functioning and services such as nutrient cycling because of increased complementarity. We examined N canopy budgets of 27 Central European forests of varying dominant tree species, stand density, and tree and shrub species diversity (Shannon index) in three study regions by quantifying bulk and fine particulate dry deposition and dissolved below canopy N fluxes. Average regional canopy N retention ranged from 16% to 51%, because of differences in the N status of the ecosystems. Canopy N budgets of coniferous forests differed from deciduous forest which we attribute to differences in biogeochemical N cycling, tree functional traits and canopy surface area. The canopy budgets of N were related to the Shannon index which explained 14% of the variance of the canopy budgets of N, suggesting complementary aboveground N use of trees and diverse understorey vegetation. The relationship between plant diversity and canopy N retention varied among regional site conditions and forest types. Our results suggest that the traditional view of belowground complementarity of nutrient uptake by roots in diverse plant communities can be transferred to foliar uptake in forest canopies.

Persistence of Artemisia steppe in the Tangra Yumco Basin, west-central Tibet, China: despite or in consequence of Holocene lake-level changes?

The closed Tangra Yumco Basin underwent the strongest Quaternary lake-level changes so far recorded on the Tibetan Plateau. It was hitherto unknown what effect this had on local Holocene vegetation development. A 3.6-m sediment core from a recessional lake terrace at 4,700 m a.s.l., 160 m above the present lake level of Tangra Yumco, was studied to reconstruct Holocene flooding phases (sedimentology and ostracod analyses), vegetation dynamics and human influence (palynology, charcoal and coprophilous fungi analyses). Peat at the base of the profile proves lake level was below 4,700 m a.s.l. during the Pleistocene/Holocene transition. A deep-lake phase started after 11 cal ka BP, but the ostracod record indicates the level was not higher than similar to 4,720 m a.s.l. (180 m above present) and decreased gradually after the early Holocene maximum. Additional sediment ages from the basin suggest recession of Tangra Yumco from the coring site after 2.6 cal ka BP, with a shallow local lake persisting at the site until similar to 1 cal ka BP. The final peat formation indicates drier conditions thereafter. Persistence of Artemisia steppe during the Holocene lake high-stand resembles palynological records from west Tibet that indicate early Holocene aridity, in spite of high lake levels that may have resulted from meltwater input. Yet pollen assemblages indicate humidity closer to that of present potential forest areas near Lhasa, with 500-600 mm annual precipitation. Thus, the early mid-Holocene humidity was sufficient to sustain at least juniper forest, but Artemisia dominance persisted as a consequence of a combination of environmental disturbances such as (1) strong early Holocene climate fluctuations, (2) inundation of habitats suitable for forest, (3) extensive water surfaces that served as barriers to terrestrial diaspore transport from refuge areas, (4) strong erosion that denuded the non-flooded upper slopes and (5) increasing human influence since the late glacial.

Phosphorus and Nitrogen Regulate Arbuscular Mycorrhizal Symbiosis in Petunia hybrida

Phosphorus and nitrogen are essential nutrient elements that are needed by plants in large amounts. The arbuscular mycorrhizal symbiosis between plants and soil fungi improves phosphorus and nitrogen acquisition under limiting conditions. On the other hand, these nutrients influence root colonization by mycorrhizal fungi and symbiotic functioning. This represents a feedback mechanism that allows plants to control the fungal symbiont depending on nutrient requirements and supply. Elevated phosphorus supply has previously been shown to exert strong inhibition of arbuscular mycorrhizal development. Here, we address to what extent inhibition by phosphorus is influenced by other nutritional pathways in the interaction between Petunia hybrida and R. irregularis. We show that phosphorus and nitrogen are the major nutritional determinants of the interaction. Interestingly, the symbiosis-promoting effect of nitrogen starvation dominantly overruled the suppressive effect of high phosphorus nutrition onto arbuscular mycorrhiza, suggesting that plants promote the symbiosis as long as they are limited by one of the two major nutrients. Our results also show that in a given pair of symbiotic partners (Petunia hybrida and R. irregularis), the entire range from mutually symbiotic to parasitic can be observed depending on the nutritional conditions. Taken together, these results reveal complex nutritional feedback mechanisms in the control of root colonization by arbuscular mycorrhizal fungi.

Placing unprecedented recent fir growth in a European-wide and Holocene-long context

Forest decline played a pivotal role in motivating Europe's political focus on sustainability around 35 years ago. Silver fir (Abies alba) exhibited a particularly severe dieback in the mid-1970s, but disentangling biotic from abiotic drivers remained challenging because both spatial and temporal data were lacking. Here, we analyze 14 136 samples from living trees and historical timbers, together with 356 pollen records, to evaluate recent fir growth from a continent-wide and Holocene-long perspective. Land use and climate change influenced forest growth over the past millennium, whereas anthropogenic emissions of acidic sulfates and nitrates became important after about 1850. Pollution control since the 1980s, together with a warmer but not drier climate, has facilitated an unprecedented surge in productivity across Central European fir stands. Restricted fir distribution prior to the Mesolithic and again in the Modern Era, separated by a peak in abundance during the Bronze Age, is indicative of the long-term interplay of changing temperatures, shifts in the hydrological cycle, and human impacts that have shaped forest structure and productivity.

Reprint of "Seed availability in hay meadows: Land-use intensification promotes seed rain but not the persistent seed bank''

Intensification of land use in semi-natural hay meadows has resulted in a decrease in species diversity. This is often thought to be caused by the reduced establishment of plant species due to high competition for light under conditions of increased productivity. Sowing experiments in grasslands have found reliable evidence that diversity can also be constrained by seed availability, implying that processes influencing the production and persistence of seeds may be important for the functioning of ecosystems. So far, the effects of land-use intensification on the seed rain and the persistence of seeds in the soil have been unclear. We selected six pairs of extensively managed (Festuco-Brometea) and intensively managed (Arrhenatheretalia) grassland with traditional late cutting regimes across Switzerland and covering an annual productivity gradient in the range 176-1211 gm(-2). In each grassland community, we estimated seed rain and seed bank using eight pooled seed-trap or topsoil samples of 89 cm(2) in each of six plots representing an area of c. 150 m(2). The seed traps were established in spring 2010 and collected simultaneously with soil cores after an exposure of c. three months. We applied the emergence method in a cold frame over eight months to estimate density of viable seeds. With community productivity reflecting land-use intensification, the density and species richness in the seed rain increased, while mean seed size diminished and the proportions of persistent seeds and of species with persistent seeds in the topsoil declined. Stronger limitation of seeds in extensively managed semi-natural grasslands can explain the fact that such grasslands are not always richer in species than more intensively managed ones. (C) 2013 Elsevier B.V. All rights reserved.

Identification and quantification of phytochelatins in roots of rice to long-term exposure: evidence of individual role on arsenic accumulation and translocation

Rice has the predilection to take up arsenic in the form of methylated arsenic (o-As) and inorganic arsenic species (i-As). Plants defend themselves using i-As efflux systems and the production of phytochelatins (PCs) to complex i-As. Our study focused on the identification and quantification of phytochelatins by HPLC-ICP-MS/ESI-MS, relating them to the several variables linked to As exposure. GSH, 11 PCs, and As–PC complexes from the roots of six rice cultivars (Italica Carolina, Dom Sofid, 9524, Kitrana 508, YRL-1, and Lemont) exposed to low and high levels of i-As were compared with total, i-As, and o-As in roots, shoots, and grains. Only Dom Sofid, Kitrana 508, and 9524 were found to produce higher levels of PCs even when exposed to low levels of As. PCs were only correlated to i-As in the roots (r=0.884, P <0.001). However, significant negative correlations to As transfer factors (TF) roots–grains (r= –0.739, P <0.05) and shoots–grains (r= –0.541, P <0.05), suggested that these peptides help in trapping i-As but not o-As in the roots, reducing grains’ i-As. Italica Carolina reduced i-As in grains after high exposure, where some specific PCs had a special role in this reduction. In Lemont, exposure to elevated levels of i-As did not result in higher i-As levels in the grains and there were no significant increases in PCs or thiols. Finally, the high production of PCs in Kitrana 508 and Dom Sofid in response to high As treatment did not relate to a reduction of i-As in grains, suggesting that other mechanisms such as As–PC release and transport seems to be important in determining grain As in these cultivars.

The effect of abrupt climatic warming on biogeochemical cycling and N2O emissions in a terrestrial ecosystem

The large, rapid increase in atmospheric N2O concentrations that occurred concurrent with the abrupt warming at the end of the Last Glacial period might have been the result of a reorganization in global biogeochemical cycles. To explore the sensitivity of nitrogen cycling in terrestrial ecosystems to abrupt warming, we combined a scenario of climate and vegetation composition change based on multiproxy data for the Oldest Dryas–Bølling abrupt warming event at Gerzensee, Switzerland, with a biogeochemical model that simulates terrestrial N uptake and release, including N2O emissions. As for many central European sites, the pollen record at the Gerzensee is remarkable for the abundant presence of the symbiotic nitrogen fixer Hippophaë rhamnoides (L.) during the abrupt warming that also marks the beginning of primary succession on immature glacial soils. Here we show that without additional nitrogen fixation, climate change results in a significant increase of N2O emissions of approximately factor 3.4 (from 6.4 ± 1.9 to 21.6 ± 5.9 mg N2O–N m− 2 yr− 1). Each additional 1000 mg m− 2 yr− 1 of nitrogen added to the ecosystem through N-fixation results in additional N2O emissions of 1.6 mg N2O–N m− 2 yr− 1 for the time with maximum H. rhamnoides coverage. Our results suggest that local reactions of emissions to abrupt climate change could have been considerably faster than the overall atmospheric concentration changes observed in polar ice. Nitrogen enrichment of soils due to the presence of symbiotic N-fixers during early primary succession not only facilitates the establishment of vegetation on soils in their initial stage of development, but can also have considerable influence on biogeochemical cycles and the release of reactive nitrogen trace gases to the atmosphere.