Drivers of biogenic volatile organic compound emissions in hygrophytic bryophytes.

Bryophytes can both emit and take up biogenic volatile organic compounds (BVOCs) to and from the environment. Despite the scarce study of these exchanges, BVOCs have been shown to be important for a wide range of ecological roles. Bryophytes are the most ancient clade of land plants and preserve very similar traits to those first land colonisers. Therefore, the study of these plants can help understand the early processes of BVOC emissions as an adaptation to terrestrial life. Here, we determine the emission rates of BVOCs from different bryophyte species to understand what drives such emissions. We studied 26 bryophyte species from temperate regions that can be found in mountain springs located in NE Spain. Bryophyte BVOC emission presented no significant phylogenetic signal for any of the compounds analysed. Hence, we used mixed linear models to investigate the species-specific differences and eco-physiological and environmental drivers of bryophyte BVOC emission. In general, species-specific variability was the main factor explaining bryophyte BVOC emissions; but additionally, photosynthetic rates and light intensity increased BVOC emissions. Despite emission measurements reported here were conducted at 30°, and may not directly correspond to emission rates in natural conditions, most of the screened species have never been measured before for BVOC emissions and therefore this information can help understand the drivers of the emissions of BVOCs in bryophytes.

Fertile forests produce biomass more efficiently.

Trees with sufficient nutrition are known to allocate carbon preferentially to aboveground plant parts. Our global study of 49 forests revealed an even more fundamental carbon allocation response to nutrient availability: forests with high-nutrient availability use 58 ± 3% (mean ± SE; 17 forests) of their photosynthates for plant biomass production (BP), while forests with low-nutrient availability only convert 42 ± 2% (mean ± SE; 19 forests) of annual photosynthates to biomass. This nutrient effect largely overshadows previously observed differences in carbon allocation patterns among climate zones, forest types and age classes. If forests with low-nutrient availability use 16 ± 4% less of their photosynthates for plant growth, what are these used for? Current knowledge suggests that lower BP per unit photosynthesis in forests with low- versus forests with high-nutrient availability reflects not merely an increase in plant respiration, but likely results from reduced carbon allocation to unaccounted components of net primary production, particularly root symbionts.

Insights into ozone deposition patterns from decade-long ozone flux measurements over a mixed temperate forest.

Long-term fluxes of ozone (O(3)) were measured over a mixed temperate forest using the aerodynamic gradient method. The long-term average O(3) flux (F) was -366 ng m(-2) s(-1) for the period 2000-2010, corresponding to an average O(3) concentration of 48 µg m(-3) and a deposition velocity v(d) of 9 mm s(-1). Average nocturnal ozone deposition amounted to -190 ng m(-2) s(-1), which was about one third of the daytime flux. Also during the winter period substantial O(3) deposition was measured. In addition, total O(3) fluxes were found to differ significantly among canopy wetness categories. During the day, highest deposition fluxes were generally measured for a dry canopy, whereas a rain-wetted canopy constituted the best sink at night. Flux partitioning calculations revealed that the stomatal flux (F(s)) contributed 20% to the total F but the F(s)/F fraction was subject to seasonal and diurnal changes. The annual concentration-based index AOT40 (accumulated dose over a threshold of 40 ppb) and the Phytotoxic Ozone Dose (POD(1) or accumulated stomatal flux above a threshold of 1 nmol m(-2) s(-1)) were related in a curvilinear way. The O(3) deposition was found to be largely controlled by non-stomatal sinks, whose strength was enhanced by high friction velocities (u(*)), optimizing the mechanical mixing of O(3) into the canopy and the trunk space. The long-term geometrical mean of the non-stomatal resistance (R(ns)) was 136 s m(-1) but lower R(ns) values were encountered during the winter half-year due to higher u(*). The R(ns) was also subject to a marked diurnal variability, with low R(ns) in the morning hours, when turbulence took off. We speculate that non-stomatal deposition was largely driven by scavenging of ozone by biogenic volatile organic compounds (BVOCs) and especially NO emitted from the crown or the forest floor.

Soil [N] modulates soil C cycling in CO2-fumigated tree stands: a meta-analysis.

Under elevated atmospheric CO(2) concentrations, soil carbon (C) inputs are typically enhanced, suggesting larger soil C sequestration potential. However, soil C losses also increase and progressive nitrogen (N) limitation to plant growth may reduce the CO(2) effect on soil C inputs with time. We compiled a data set from 131 manipulation experiments, and used meta-analysis to test the hypotheses that: (1) elevated atmospheric CO(2) stimulates soil C inputs more than C losses, resulting in increasing soil C stocks; and (2) that these responses are modulated by N. Our results confirm that elevated CO(2) induces a C allocation shift towards below-ground biomass compartments. However, the increased soil C inputs were offset by increased heterotrophic respiration (Rh), such that soil C content was not affected by elevated CO(2). Soil N concentration strongly interacted with CO(2) fumigation: the effect of elevated CO(2) on fine root biomass and -production and on microbial activity increased with increasing soil N concentration, while the effect on soil C content decreased with increasing soil N concentration. These results suggest that both plant growth and microbial activity responses to elevated CO(2) are modulated by N availability, and that it is essential to account for soil N concentration in C cycling analyses.

Nutrient scarcity as a selective pressure for mast seeding.

Mast seeding is one of the most intriguing reproductive traits in nature. Despite its potential drawbacks in terms of fitness, the widespread existence of this phenomenon suggests that it should have evolutionary advantages under certain circumstances. Using a global dataset of seed production time series for 219 plant species from all of the continents, we tested whether masting behaviour appears predominantly in species with low foliar nitrogen and phosphorus concentrations when controlling for local climate and productivity. Here, we show that masting intensity is higher in species with low foliar N and P concentrations, and especially in those with imbalanced N/P ratios, and that the evolutionary history of masting behaviour has been linked to that of nutrient economy. Our results support the hypothesis that masting is stronger in species growing under limiting conditions and suggest that this reproductive behaviour might have evolved as an adaptation to nutrient limitations and imbalances.

Global patterns of phosphatase activity in natural soils.

Soil phosphatase levels strongly control the biotic pathways of phosphorus (P), an essential element for life, which is often limiting in terrestrial ecosystems. We investigated the influence of climatic and soil traits on phosphatase activity in terrestrial systems using metadata analysis from published studies. This is the first analysis of global measurements of phosphatase in natural soils. Our results suggest that organic P (Porg), rather than available P, is the most important P fraction in predicting phosphatase activity. Structural equation modeling using soil total nitrogen (TN), mean annual precipitation, mean annual temperature, thermal amplitude and total soil carbon as most available predictor variables explained up to 50% of the spatial variance in phosphatase activity. In this analysis, Porg could not be tested and among the rest of available variables, TN was the most important factor explaining the observed spatial gradients in phosphatase activity. On the other hand, phosphatase activity was also found to be associated with climatic conditions and soil type across different biomes worldwide. The close association among different predictors like Porg, TN and precipitation suggest that P recycling is driven by a broad scale pattern of ecosystem productivity capacity.

Atmospheric deposition, CO2, and change in the land carbon sink.

Concentrations of atmospheric carbon dioxide (CO2) have continued to increase whereas atmospheric deposition of sulphur and nitrogen has declined in Europe and the USA during recent decades. Using time series of flux observations from 23 forests distributed throughout Europe and the USA, and generalised mixed models, we found that forest-level net ecosystem production and gross primary production have increased by 1% annually from 1995 to 2011. Statistical models indicated that increasing atmospheric CO2 was the most important factor driving the increasing strength of carbon sinks in these forests. We also found that the reduction of sulphur deposition in Europe and the USA lead to higher recovery in ecosystem respiration than in gross primary production, thus limiting the increase of carbon sequestration. By contrast, trends in climate and nitrogen deposition did not significantly contribute to changing carbon fluxes during the studied period. Our findings support the hypothesis of a general CO2-fertilization effect on vegetation growth and suggest that, so far unknown, sulphur deposition plays a significant role in the carbon balance of forests in industrialized regions. Our results show the need to include the effects of changing atmospheric composition, beyond CO2, to assess future dynamics of carbon-climate feedbacks not currently considered in earth system/climate modelling.

Evaluating the convergence between eddy-covariance and biometric methods for assessing carbon budgets of forests.

The eddy-covariance (EC) micro-meteorological technique and the ecology-based biometric methods (BM) are the primary methodologies to quantify CO2 exchange between terrestrial ecosystems and the atmosphere (net ecosystem production, NEP) and its two components, ecosystem respiration and gross primary production. Here we show that EC and BM provide different estimates of NEP, but comparable ecosystem respiration and gross primary production for forest ecosystems globally. Discrepancies between methods are not related to environmental or stand variables, but are consistently more pronounced for boreal forests where carbon fluxes are smaller. BM estimates are prone to underestimation of net primary production and overestimation of leaf respiration. EC biases are not apparent across sites, suggesting the effectiveness of standard post-processing procedures. Our results increase confidence in EC, show in which conditions EC and BM estimates can be integrated, and which methodological aspects can improve the convergence between EC and BM.

Contrasting net primary productivity and carbon distribution between neighboring stands of Quercus robur and Pinus sylvestris.

Standing biomass, net primary production (NPP) and soil carbon (C) pools were studied in a 67-year-old pedunculate oak (Quercus robur L.) stand and a neighboring 74-year- old Scots pine (Pinus sylvestris L.) stand in the Belgian Campine region. Despite a 14% lower tree density and a lower tree height in the oak stand, standing biomass was slightly higher than in the pine stand (177 and 169 Mg ha(-1) in oaks and pines, respectively), indicating that individual oak trees contained more biomass than pine trees of similar diameter. Moreover, NPP in the oak stand was more than double that in the pine stand (17.7 and 8.1 Mg ha(-1) year(-1), respectively). Several observations indicated that soil organic matter accumulated at higher rates under pines than under oaks. We therefore hypothesized that the pines were exhibiting an age-related decline in productivity due to nutrient limitation. The poor decomposability of pine litter resulted in the observed accumulation of organic matter. The subsequent immobilization of nutrients in the organic matter, combined with the already nutrient-poor soil conditions, resulted in a decrease in total NPP over time, as well as in a substantial shift in the allocation of NPP toward fine roots. In the oak stand, litter is less recalcitrant to decay and soil acidity is less severe; hence, organic matter does not accumulate and nutrients are recycled. This probably explains why NPP was much higher in the oaks than in the pines and why only a small proportion of NPP was allocated to oak fine roots.

Soil respiration in a mixed temperate forest and its contribution to total ecosystem respiration.

Soil respiration (SR) was measured with an infrared gas analyzer in nine plots representative of the heterogeneous vegetation in a mixed coniferous-deciduous forest in the Belgian Campine region. Selected plots included the two most representative overstory species (Pinus sylvestris L. and Quercus robur L.) in combination with the most representative understory species of the forest. A model that includes temperature and water as the main controlling variables was fitted to the data. We found large spatial variability in SR among plots, with typically lower fluxes under the coniferous overstory than under the deciduous overstory (means of 4.8 +/- 0.4 and 8.8 +/- 0.5 Mg C ha(-1) year(-1), respectively). Total annual soil carbon (C) emissions were estimated by weighting fluxes from different types of vegetation according to their relative contribution to the footprint area of the eddy covariance flux measurement. The relative contribution of the two main tree species to the footprint-weighted total SR varied among seasons with the more abundant coniferous overstory contributing the most to total SR during most of the year. Nonetheless, during summer, the contribution of deciduous plots to total SR was disproportionally high because of the more pronounced seasonality of belowground metabolic activity. Net ecosystem carbon dioxide exchange was measured by eddy covariance, and we estimated total ecosystem respiration (TER) with footprint-constrained nighttime fluxes. Mean total annual SR and TER were 6.1 +/- 0.11 and 9.1 +/- 1.15 Mg C ha(-1) year(-1), respectively. The 95% confidence interval of the ratio of annual SR:TER ranged from 0.58 to 0.76, with a mean of 0.67. The contribution of SR to TER tended to vary seasonally, with minimum contributions during summer (less than 50% of TER) and maximum contributions during winter (about 94% of TER).

Net carbon storage in a poplar plantation (POPFACE) after three years of free-air CO2 enrichment.

A high-density plantation of three genotypes of Populus was exposed to an elevated concentration of carbon dioxide ([CO(2)]; 550 micromol mol(-1)) from planting through canopy closure using a free-air CO(2) enrichment (FACE) technique. The FACE treatment stimulated gross primary productivity by 22 and 11% in the second and third years, respectively. Partitioning of extra carbon (C) among C pools of different turnover rates is of critical interest; thus, we calculated net ecosystem productivity (NEP) to determine whether elevated atmospheric [CO(2)] will enhance net plantation C storage capacity. Free-air CO(2) enrichment increased net primary productivity (NPP) of all genotypes by 21% in the second year and by 26% in the third year, mainly because of an increase in the size of C pools with relatively slow turnover rates (i.e., wood). In all genotypes in the FACE treatment, more new soil C was added to the total soil C pool compared with the control treatment. However, more old soil C loss was observed in the FACE treatment compared with the control treatment, possibly due to a priming effect from newly incorporated root litter. FACE did not significantly increase NEP, probably as a result of this priming effect.

Interactive effects of temperature and precipitation on soil respiration in a temperate maritime pine forest.

Soil respiration (SR) was monitored periodically throughout 2001 in a Scots pine (Pinus sylvestris L.) stand located in the Belgian Campine region. As expected for a temperate maritime forest, temperature was the dominant control over SR during most of the year. However, during late spring and summer, when soil water content (SWC) was limiting, SR was insensitive to temperature (Q(10) = 1.24). We observed that during prolonged rain-free periods, when SWC was less than 15% (v/v), SR decreased dramatically (up to 50%) and SWC took over control of SR. During such drought periods, however, rain events sometimes stimulated SR and restored temperature control over SR, even though SWC in the mineral soil was low. We hypothesize that restoration of temperature control occurred only when rain events adequately rewetted the uppermost soil layers, where most of the respiratory activity occurred. To quantify the rewetting capacity of rain events, an index (I(w)) was designed that incorporated rainfall intensity, time elapsed since the last rain event, and atmospheric vapor pressure deficit (a proxy for evaporative water losses). To simulate SR fluxes, a model was developed that included the effects of soil temperature and, under drought and non-rewetting conditions (I(w) and SWC < threshold), an SWC response function. The model explained 95% of the temporal variability in SR observed during summer, whereas the temperature function alone explained only 73% of this variability. Our results revealed that, in addition to temperature and SWC, rain plays a role in determining the total amount of carbon released from soils, even in a maritime climate.

Above- and belowground biomass and net primary production in a 73-year-old Scots pine forest.

We estimated above- and belowground biomass and net primary production (NPP) of a 73-year-old Scots pine (Pinus sylvestris L.) forest stand in the Belgian Campine region. Total biomass for the stand was 176 Mg ha(-1), of which 74.4% was found in stems. The root system contained 12.6% of total biomass, most of it in coarse roots (> 5 mm). Fine roots (< 5 mm) comprised only about 1.7% of total biomass, and more than 50% of fine root biomass was retrieved in the litter layer and the upper 15 cm of the mineral soil. The ratio of belowground biomass to aboveground biomass was 0.14, which is lower than that of other Scots pine forests and other coniferous forests. Between 1995 and 2001, mean annual NPP was 11.2 Mg ha(-1) year(-1), of which 68.7% was allocated to aboveground compartments. Stems, needles and cones made relatively high contributions to total NPP compared with branches. However, branch NPP was possibly underestimated because litterfall of big branches was neglected. The proportion of total NPP in belowground components was 31.3%. Coarse root NPP (2% of total) was low compared with its biomass. Fine root NPP was 3.3 Mg ha(-1) year(-1), representing about 29.5% of total NPP; however, the estimate of fine root NPP is much more uncertain than NPP of aboveground compartments. The ratio NPP/GPP (gross primary production) was 0.32, which was low compared with other coniferous forests.

Fine root biomass and turnover of two fast-growing poplar genotypes in a short-rotation coppice culture.

BACKGROUND AND AIMS: The quantification of root dynamics remains a major challenge in ecological research because root sampling is laborious and prone to error due to unavoidable disturbance of the delicate soil-root interface. The objective of the present study was to quantify the distribution of the biomass and turnover of roots of poplars (Populus) and associated understory vegetation during the second growing season of a high-density short rotation coppice culture. METHODS: Roots were manually picked from soil samples collected with a soil core from narrow (75 cm apart) and wide rows (150 cm apart) of the double-row planting system from two genetically contrasting poplar genotypes. Several methods of estimating root production and turnover were compared. RESULTS: Poplar fine root biomass was higher in the narrow rows than in the wide rows. In spite of genetic differences in above-ground biomass, annual fine root productivity was similar for both genotypes (ca. 44 g DM m-2 year-1). Weed root biomass was equally distributed over the ground surface, and root productivity was more than two times higher compared to poplar fine roots (ca. 109 g DM m-2 year-1). CONCLUSIONS: Early in SRC plantation development, weeds result in significant root competition to the crop tree poplars, but may confer certain ecosystem services such as carbon input to soil and retention of available soil N until the trees fully occupy the site.

Soil respiration at mean annual temperature predicts annual total across vegetation types and biomes.

Soil respiration (SR) constitutes the largest flux of CO(2) from terrestrial ecosystems to the atmosphere. However, there still exist considerable uncertainties as to its actual magnitude, as well as its spatial and interannual variability. Based on a reanalysis and synthesis of 80 site-years for 57 forests, plantations, savannas, shrublands and grasslands from boreal to tropical climates we present evidence that total annual SR is closely related to SR at mean annual soil temperature (SR(MAT)), irrespective of the type of ecosystem and biome. This is theoretically expected for non water-limited ecosystems within most of the globally occurring range of annual temperature variability and sensitivity (Q(10)). We further show that for seasonally dry sites where annual precipitation (P) is lower than potential evapotranspiration (PET), annual SR can be predicted from wet season SR(MAT) corrected for a factor related to P/PET. Our finding indicates that it can be sufficient to measure SR(MAT) for obtaining a well constrained estimate of its annual total. This should substantially increase our capacity for assessing the spatial distribution of soil CO(2) emissions across ecosystems, landscapes and regions, and thereby contribute to improving the spatial resolution of a major component of the global carbon cycle.

Temperature dependence of greenhouse gas emissions from three hydromorphic soils at different groundwater levels.

Wetlands contribute considerably to the global greenhouse gas (GHG) balance. In these ecosystems, groundwater level (GWL) and temperature, two factors likely to be altered by climate change, exert important control over CO(2), CH(4) and N(2)O fluxes. However, little is known about the temperature sensitivity (Q(10)) of the combined GHG emissions from hydromorphic soils and how this Q(10) varies with GWL. We performed a greenhouse experiment in which three different (plant-free) hydromorphic soils from a temperate spruce forest were exposed to two GWLs (an intermediate GWL of -20 cm and a high GWL of -5 cm). Net CO(2), CH(4) and N(2)O fluxes were measured continuously. Here, we discuss how these fluxes responded to synoptic temperature fluctuations. Across all soils and GWLs, CO(2) emissions responded similarly to temperature and Q(10) was close to 2. The Q(10) of the CH(4) and N(2)O fluxes also was similar across soil types. GWL, on the other hand, significantly affected the Q(10) of both CH(4) and N(2)O emissions. The Q(10) of the net CH(4) fluxes increased from about 1 at GWL = -20 cm to 3 at GWL = -5 cm. For the N(2)O emissions, Q(10) varied around 2 for GWL = -20 cm and around 4 for GWL = -5 cm. This substantial GWL-effect on the Q(10) of CH(4) and N(2)O emissions was, however, hardly reflected in the Q(10) of the total GHG emissions (which varied around 2), because the contribution of these gases was relatively small compared to that of CO(2).