Soil evaporation and organic matter turnover in the Sub-Taiga and Forest-Steppe of southwest Siberia.

Southwest Siberia encompasses the forest-steppe and sub-taiga climatic zones and has historically been utilized for agriculture. Coinciding with predicted changes in climate for the region is the pressure of agricultural development; however, a characterization of the soil water and carbon dynamics is lacking. We assessed current soil water properties and soil organic carbon turnover in forests and grasslands for two sites that span the forest steppe and sub-taiga bioclimatic zones. Soil evaporation was 0.62 ± 0.17 mm d-1 (mean ± standard error) in grasslands and 0.45 ± 0.08 mm d-1 in the forests of the forest-steppe site. Evaporation at the sub-taiga site was 1.80 ± 1.70 mm d-1 in grasslands and 0.96 ± 0.05 mm d-1 in forest plots. Evaporation was significantly greater at the sub-taiga site than the forest-steppe site. The density of fine roots explained the soil water isotopic patterns between vegetation types and sites. We found soil organic matter turnover to be three times faster in the sub-taiga site than in the forest-steppe site. Our results show that while climate factors, in particular snow levels, between the two sites are drivers for water and carbon cycles, site level hydrology, soil characteristics, and vegetation directly interact to influence the water and carbon dynamics.

On the contributions of photorespiration and compartmentation to the contrasting intramolecular 2H profiles of C3 and C4 plant sugars.

Compartmentation of C4 photosynthetic biochemistry into bundle sheath (BS) and mesophyll (M) cells, and photorespiration in C3 plants is predicted to have hydrogen isotopic consequences for metabolites at both molecular and site-specific levels. Molecular-level evidence was recently reported (Zhou et al., 2016), but evidence at the site-specific level is still lacking. We propose that such evidence exists in the contrasting 2H distribution profiles of glucose samples from naturally grown C3, C4 and CAM plants: photorespiration contributes to the relative 2H enrichment in H5 and relative 2H depletion in H1 & H6 (the average of the two pro-chiral Hs and in particular H6,pro-R) in C3 glucose, while 2H-enriched C3 mesophyll cellular (chloroplastic) water most likely contributes to the enrichment at H4; export of (transferable hydrogen atoms of) NADPH from C4 mesophyll cells to bundle sheath cells (via the malate shuttle) and incorporation of 2H-relatively unenriched BS cellular water contribute to the relative depletion of H4 & H5 respectively; shuttling of triose-phosphates (PGA: phosphoglycerate dand DHAP: dihydroacetone phosphate) between C4 bundle sheath and mesophyll cells contributes to the relative enrichment in H1 & H6 (in particular H6,pro-R) in C4 glucose.

Forest understory plant and soil microbial response to an experimentally induced drought and heat-pulse event: the importance of maintaining the continuum.

Drought duration and intensity are expected to increase with global climate change. How changes in water availability and temperature affect the combined plant-soil-microorganism response remains uncertain. We excavated soil monoliths from a beech (Fagus sylvatica L.) forest, thus keeping the understory plant-microbe communities intact, imposed an extreme climate event, consisting of drought and/or a single heat-pulse event, and followed microbial community dynamics over a time period of 28 days. During the treatment, we labeled the canopy with (13) CO2 with the goal of (i) determining the strength of plant-microbe carbon linkages under control, drought, heat and heat-drought treatments and (ii) characterizing microbial groups that are tightly linked to the plant-soil carbon continuum based on (13) C-labeled PLFAs. Additionally, we used 16S rRNA sequencing of bacteria from the Ah horizon to determine the short-term changes in the active microbial community. The treatments did not sever within-plant transport over the experiment, and carbon sinks belowground were still active. Based on the relative distribution of labeled carbon to roots and microbial PLFAs, we determined that soil microbes appear to have a stronger carbon sink strength during environmental stress. High-throughput sequencing of the 16S rRNA revealed multiple trajectories in microbial community shifts within the different treatments. Heat in combination with drought had a clear negative effect on microbial diversity and resulted in a distinct shift in the microbial community structure that also corresponded to the lowest level of label found in the PLFAs. Hence, the strongest changes in microbial abundances occurred in the heat-drought treatment where plants were most severely affected. Our study suggests that many of the shifts in the microbial communities that we might expect from extreme environmental stress will result from the plant-soil-microbial dynamics rather than from direct effects of drought and heat on soil microbes alone.

Water level changes affect carbon turnover and microbial community composition in lake sediments.

Due to climate change, many lakes in Europe will be subject to higher variability of hydrological characteristics in their littoral zones. These different hydrological regimes might affect the use of allochthonous and autochthonous carbon sources. We used sandy sediment microcosms to examine the effects of different hydrological regimes (wet, desiccating, and wet-desiccation cycles) on carbon turnover. (13)C-labelled particulate organic carbon was used to trace and estimate carbon uptake into bacterial biomass (via phospholipid fatty acids) and respiration. Microbial community changes were monitored by combining DNA- and RNA-based real-time PCR quantification and terminal restriction fragment length polymorphism (T-RFLP) analysis of 16S rRNA. The shifting hydrological regimes in the sediment primarily caused two linked microbial effects: changes in the use of available organic carbon and community composition changes. Drying sediments yielded the highest CO2 emission rates, whereas hydrological shifts increased the uptake of allochthonous organic carbon for respiration. T-RFLP patterns demonstrated that only the most extreme hydrological changes induced a significant shift in the active and total bacterial communities. As current scenarios of climate change predict an increase of drought events, frequent variations of the hydrological regimes of many lake littoral zones in central Europe are anticipated. Based on the results of our study, this phenomenon may increase the intensity and amplitude in rates of allochthonous organic carbon uptake and CO2 emissions.

Soil bacterial community structure responses to precipitation reduction and forest management in forest ecosystems across Germany.

Soil microbial communities play an important role in forest ecosystem functioning, but how climate change will affect the community composition and consequently bacterial functions is poorly understood. We assessed the effects of reduced precipitation with the aim of simulating realistic future drought conditions for one growing season on the bacterial community and its relation to soil properties and forest management. We manipulated precipitation in beech and conifer forest plots managed at different levels of intensity in three different regions across Germany. The precipitation reduction decreased soil water content across the growing season by between 2 to 8% depending on plot and region. T-RFLP analysis and pyrosequencing of the 16S rRNA gene were used to study the total soil bacterial community and its active members after six months of precipitation reduction. The effect of reduced precipitation on the total bacterial community structure was negligible while significant effects could be observed for the active bacteria. However, the effect was secondary to the stronger influence of specific soil characteristics across the three regions and management selection of overstorey tree species and their respective understorey vegetation. The impact of reduced precipitation differed between the studied plots; however, we could not determine the particular parameters being able to modify the response of the active bacterial community among plots. We conclude that the moderate drought induced by the precipitation manipulation treatment started to affect the active but not the total bacterial community, which points to an adequate resistance of the soil microbial system over one growing season.

Allocate carbon for a reason: priorities are reflected in the ¹³C/¹²C ratios of plant lipids synthesized via three independent biosynthetic pathways.

It has long been theorized that carbon allocation, in addition to the carbon source and to kinetic isotopic effects associated with a particular lipid biosynthetic pathway, plays an important role in shaping the carbon isotopic composition ((13)C/(12)C) of lipids (Park and Epstein, 1961). If the latter two factors are properly constrained, valuable information about carbon allocation during lipid biosynthesis can be obtained from carbon isotope measurements. Published work of Chikaraishi et al. (2004) showed that leaf lipids isotopic shifts from bulk leaf tissue Δδ(13)C(bk-lp) (defined as δ(13)C(bulkleaftissue)-δ(13)C(lipid)) are pathway dependent: the acetogenic (ACT) pathway synthesizing fatty lipids has the largest isotopic shift, the mevalonic acid (MVA) pathway synthesizing sterols the lowest and the phytol synthesizing 1-deoxy-D-xylulose 5-phosphate (DXP) pathway gives intermediate values. The differences in Δδ(13)C(bk-lp) between C3 and C4 plants Δδ(13)C(bk-lp,C4-C3) are also pathway-dependent: Δδ(13)C(ACT)(bk-lp,C4-C3) > Δδ(13)C(DXP(bk-lp,C4-C3) > Δδ(13)C(MVA)(bk-lp,C4-C3). These pathway-dependent differences have been interpreted as resulting from kinetic isotopic effect differences of key but unspecified biochemical reactions involved in lipids biosynthesis between C3 and C4 plants. After quantitatively considering isotopic shifts caused by (dark) respiration, export-of-carbon (to sink tissues) and photorespiration, we propose that the pathway-specific differences Δδ(13)C(bk-lp,C4-C3) can be successfully explained by C4-C3 carbon allocation (flux) differences with greatest flux into the ACT pathway and lowest into the MVA pathways (when flux is higher, isotopic shift relative to source is smaller). Highest carbon allocation to the ACT pathway appears to be tied to the most stringent role of water-loss-minimization by leaf waxes (composed mainly of fatty lipids) while the lowest carbon allocation to the MVA pathway can be largely explained by the fact that sterols act as regulatory hormones and membrane fluidity modulators in rather low concentrations.

The oxygen isotope enrichment of leaf-exported assimilates--does it always reflect lamina leaf water enrichment?

The oxygen stable isotope composition of plant organic matter (OM) (particularly of wood and cellulose in the tree ring archive) is valuable in studies of plant-climate interaction, but there is a lack of information on the transfer of the isotope signal from the leaf to heterotrophic tissues. We studied the oxygen isotopic composition and its enrichment above source water of leaf water over diel courses in five tree species covering a broad range of life forms. We tracked the transfer of the isotopic signal to leaf water-soluble OM and further to phloem-transported OM. Observed leaf water evaporative enrichment was consistent with values predicted from mechanistic models taking into account nonsteady-state conditions. While leaf water-soluble OM showed the expected (18)O enrichment in all species, phloem sugars were less enriched than expected from leaf water enrichment in Scots pine (Pinus sylvestris), European larch (Larix decidua) and Alpine ash (Eucalyptus delegatensis). Oxygen atom exchange with nonenriched water during phloem loading and transport, as well as a significant contribution of assimilates from bark photosynthesis, can explain these phloem (18)O enrichment patterns. Our results indicate species-specific uncoupling between the leaf water and the OM oxygen isotope signal, which is important for the interpretation of tree ring data.

Soil moisture effects on the carbon isotope composition of soil respiration.

The carbon isotopic composition (delta(13)C) of recently assimilated plant carbon is known to depend on water-stress, caused either by low soil moisture or by low atmospheric humidity. Air humidity has also been shown to correlate with the delta(13)C of soil respiration, which suggests indirectly that recently fixed photosynthates comprise a substantial component of substrates consumed by soil respiration. However, there are other reasons why the delta(13)CO(2) of soil efflux may change with moisture conditions, which have not received as much attention. Using a combination of greenhouse experiments and modeling, we examined whether moisture can cause changes in fractionation associated with (1) non-steady-state soil CO(2) transport, and (2) heterotrophic soil-respired delta(13)CO(2). In a first experiment, we examined the effects of soil moisture on total respired delta(13)CO(2) by growing Douglas fir seedlings under high and low soil moisture conditions. The measured delta(13)C of soil respiration was 4.7 per thousand more enriched in the low-moisture treatment; however, subsequent investigation with an isotopologue-based gas diffusion model suggested that this result was probably influenced by gas transport effects. A second experiment examined the heterotrophic component of soil respiration by incubating plant-free soils, and showed no change in microbial-respired delta(13)CO(2) across a large moisture range. Our results do not rule out the potential influence of recent photosynthates on soil-respired delta(13)CO(2), but they indicate that the expected impacts of photosynthetic discrimination may be similar in direction and magnitude to those from gas transport-related fractionation. Gas transport-related fractionation may operate as an alternative or an additional factor to photosynthetic discrimination to explain moisture-related variation in soil-respired delta(13)CO(2).

Bias and uncertainty of delta13CO 2 isotopic mixing models.

Patterns in the isotopic signal (stable C isotope composition; delta(13)C) of respiration (delta(13)C(R)) have led to important gains in understanding the C metabolism of many systems. Contained within delta(13)C(R) is a record of the C source mineralized, the metabolic pathway of C and the environmental conditions during which respiration occurred. Because gas samples used for analysis of delta(13)C(R) contain a mixture of CO(2) from respiration and from the atmosphere, two-component mixing models are used to identify delta(13)C(R). Measurement of ecosystem delta(13)C(R), using canopy airspace gas samples, was one of the first applications of mixing models in ecosystem ecology, and thus recommendations and guidelines are based primarily on findings from these studies. However, as mixing models are applied to other experimental conditions these approaches may not be appropriate. For example, the range in [CO(2)] obtained in gas samples from canopy air is generally less than 100 micromol mol(-1), whereas in studies of respiration from soil, foliage or tree stems, the range can span as much as 10,000 micromol mol(-1) and greater. Does this larger range in [CO(2)] influence the precision and accuracy of delta(13)C(R) estimates derived from mixing models? Does the outcome from using different regression approaches and mixing models vary depending on the range of [CO(2)]? Our research addressed these questions using a simulation approach. We found that it is important to distinguish between large (>1,000 micromol mol(-1)) and small (<100 micromol mol(-1)) ranges of CO(2) when applying a mixing model (Keeling plot or Miller-Tans) and regression approach (ordinary least squares or geometric mean regression) combination to isotopic data. The combination of geometric mean regression and the Miller-Tans mixing model provided the most accurate and precise estimate of delta(13)C(R) when the range of CO(2) is >or=1,000 micromol mol(-1).

A laboratory comparison of two methods used to estimate the isotopic composition of soil delta13CO2 efflux at steady state.

The stable isotopic composition of soil (13)CO(2) flux is important for monitoring soil biological and physical processes. While several methods exist to measure the isotopic composition of soil flux, we do not know how effective each method is at achieving this goal. To provide clear evidence of the accuracy of current measurement techniques we created a column filled with quartz sand through which a gas of known isotopic composition (-34.2 per thousand) and concentration (3,000 ppm) diffused for 7 h. We used a static chamber at equilibrium and a soil probe technique to test whether they could identify the isotopic signature of the known gas source. The static chamber is designed to identify the source gas isotopic composition when in equilibrium with the soil gas, and the soil probe method relies on a mixing model of samples withdrawn from three gas wells at different depths to identify the gas source. We sampled from ports installed along the side of the sand column to describe the isotopic and concentration gradient as well as to serve as a control for the soil probe. The soil probe produced similar isotopic and concentration values as the control ports, as well as Keeling intercepts. The static chamber at equilibrium did not identify the source gas but, when applied in a two end-member mixing model, did produce a similar Keeling intercept produced from the control ports. Neither of the methods was able to identify the source gas via the Keeling plot method probably because CO(2) profiles did not reach isotopic steady state. Our results showed that the static chamber at equilibrium should be used only with a Keeling plot approach and that the soil probe is able to provide estimates of uncertainty for the isotopic composition of soil gas as well as information pertinent to the soil profile.