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.

Drought priming reduces Calligonum mongolicum sensitivity to recurrent droughts via coordinated regulation of osmolytes, antioxidants, and hormones.

Pre-exposure of plants to abiotic stressors may induce stress memory and improve tolerance to subsequent stresses. Here, 3-month-old Calligonum mongolicum seedlings were exposed to drought (60 days) with (primed) or without (unprimed) early drought exposure of 50 days, to determine whether this enhances seedling resistance and investigate possible underlying mechanisms. Compared to unprimed, primed seedlings had higher biomass, shoot relative water content (15% and 22%), chlorophyll a, chlorophyll b, and carotenoids. They also had more superoxide anions (O2 -• ) and H2 O2 scavenging mechanisms through higher activity of SOD, CAT, APX, and dehydroascorbate reductase in assimilating shoots and roots, resulting in less ROS and oxidative stress damage. Plants also had higher ABA and JA but lower SA, likely reflecting an adaptive response to subsequent stress. Primed seedlings accumulated more IAA and brassinosteroids, which may account for their better growth. Accumulation of glycine betaine, pro, and total amino acids in assimilating shoots and roots of primed seedlings led to reduced osmotic stress. Drivers of responses of non-primed and primed seedlings to drought varied. Responses of primed seedlings were primarily characterized by more photosynthetic pigments, increased oxidative scavenging of O2 -• and H2 O2 , more phytohormones and osmolytes. Early drought priming of drought stress memory in C. mongolicum seedlings may provide a useful management approach to improve seedling establishment in vegetation restoration programs.

Mediterranean springs: Keystone ecosystems and biodiversity refugia threatened by global change.

Mediterranean spring ecosystems are unique habitats at the interface between surface water and groundwater. These ecosystems support a remarkable array of biodiversity and provide important ecological functions and ecosystem services. Spring ecosystems are influenced by abiotic, biotic, and anthropogenic factors such as the lithology of their draining aquifers, their climate, and the land use of their recharge area, all of which affect the water chemistry of the aquifer and the spring discharges. One of the most relevant characteristics of spring ecosystems is the temporal stability of environmental conditions, including physicochemical features of the spring water, across seasons and years. This stability allows a wide range of species to benefit from these ecosystems (particularly during dry periods), fostering an unusually high number of endemic species. However, global change poses important threats to these freshwater ecosystems. Changes in temperature, evapotranspiration, and precipitation patterns can alter the water balance and chemistry of spring water. Eutrophication due to agricultural practices and emergent pollutants, such as pharmaceuticals, personal care products, and pesticides, is also a growing concern for the preservation of spring biodiversity. Here, we provide a synthesis of the main characteristics and functioning of Mediterranean spring ecosystems. We then describe their ecological value and biodiversity patterns and highlight the main risks these ecosystems face. Moreover, we identify existing knowledge gaps to guide future research in order to fully uncover the hidden biodiversity within these habitats and understand the main drivers that govern them. Finally, we provide a brief summary of recommended actions that should be taken to effectively manage and preserve Mediterranean spring ecosystems for future generations. Even though studies on Mediterranean spring ecosystems are still scarce, our review shows there are sufficient data to conclude that their future viability as functional ecosystems is under severe threat.

Assessing the insulating properties of an ultrathin SrTiO3shell grown around GaAs nanowires with molecular beam epitaxy.

We have studied electronic transport in undoped GaAs/SrTiO3core-shell nanowires standing on their Si substrate with two-tip scanning tunneling microscopy in ultrahigh vacuum. The resistance profile along the nanowires is proportional to the tip separation with resistances per unit length of a few GΩ/µm. Examination of the different transport pathways parallel to the nanowire growth axis reveals that the measured resistance is consistent with a conduction along the interfacial states at the GaAs{110} sidewalls, the 2 nm thick SrTiO3shell being as much as resistive, despite oxygen deficient growth conditions. The origin of the shell resistivity is discussed in light of the nanowire analysis with transmission electron microscopy and Raman spectroscopy, providing good grounds for the use of SrTiO3shells as gate insulators.

Metabolomics and transcriptomics to decipher molecular mechanisms underlying ectomycorrhizal root colonization of an oak tree.

Mycorrhizas are known to have a positive impact on plant growth and ability to resist major biotic and abiotic stresses. However, the metabolic alterations underlying mycorrhizal symbiosis are still understudied. By using metabolomics and transcriptomics approaches, cork oak roots colonized by the ectomycorrhizal fungus Pisolithus tinctorius were compared with non-colonized roots. Results show that compounds putatively corresponding to carbohydrates, organic acids, tannins, long-chain fatty acids and monoacylglycerols, were depleted in ectomycorrhizal cork oak colonized roots. Conversely, non-proteogenic amino acids, such as gamma-aminobutyric acid (GABA), and several putative defense-related compounds, including oxylipin-family compounds, terpenoids and B6 vitamers were induced in mycorrhizal roots. Transcriptomic analysis suggests the involvement of GABA in ectomycorrhizal symbiosis through increased synthesis and inhibition of degradation in mycorrhizal roots. Results from this global metabolomics analysis suggest decreases in root metabolites which are common components of exudates, and in compounds related to root external protective layers which could facilitate plant-fungal contact and enhance symbiosis. Root metabolic pathways involved in defense against stress were induced in ectomycorrhizal roots that could be involved in a plant mechanism to avoid uncontrolled growth of the fungal symbiont in the root apoplast. Several of the identified symbiosis-specific metabolites, such as GABA, may help to understand how ectomycorrhizal fungi such as P. tinctorius benefit their host plants.

Wurtzite phase control for self-assisted GaAs nanowires grown by molecular beam epitaxy.

The accurate control of the crystal phase in III-V semiconductor nanowires (NWs) is an important milestone for device applications. Although cubic zinc-blende (ZB) GaAs is a well-established material in microelectronics, the controlled growth of hexagonal wurtzite (WZ) GaAs has thus far not been achieved successfully. Specifically, the prospect of growing defect-free and gold catalyst-free wurtzite GaAs would pave the way towards integration on silicon substrate and new device applications. In this article, we present a method to select and maintain the WZ crystal phase in self-assisted NWs by molecular beam epitaxy. By choosing a specific regime where the NW growth process is a self-regulated system, the main experimental parameter to select the ZB or WZ phase is the V/III flux ratio. Using an analytical growth model, we show that the V/III flux ratio can be finely tuned by changing the As flux, thus driving the system toward a stationary regime where the wetting angle of the Ga droplet can be maintained in the range of values allowing the formation of pure WZ phase. The analysis of the in situ reflection high energy electron diffraction evolution, combined with high-resolution scanning transmission electron microscopy (TEM), dark field TEM, and photoluminescence all confirm the control of an extended pure WZ segment, more than a micrometer long, obtained by molecular beam epitaxy growth of self- assisted GaAs NWs with a V/III flux ratio of 4.0. This successful controlled growth of WZ GaAs suggests potential benefits for electronics and opto-electronics applications.

Crystal phase engineering of self-catalyzed GaAs nanowires using a RHEED diagram.

It is well known that the crystalline structure of the III-V nanowires (NWs) is mainly controlled by the wetting contact angle of the catalyst droplet which can be tuned by the III and V flux. In this work we present a method to control the wurtzite (WZ) or zinc-blende (ZB) structure in self-catalyzed GaAs NWs grown by molecular beam epitaxy, using in situ reflection high energy electron diffraction (RHEED) diagram analysis. Since the diffraction patterns of the ZB and WZ structures differ according to the azimuth [11̄0], it is possible to follow the evolution of the intensity of specific ZB and WZ diffraction spots during NW growth as a function of the growth parameters such as the Ga flux. By analyzing the evolution of the WZ and ZB spot intensities during NW growth with specific changes of the Ga flux, it is then possible to control the crystal structure of the NWs. ZB GaAs NWs with a controlled WZ segment have thus been realized. Using a semi-empirical model for the NW growth and our in situ RHEED measurements, the critical wetting angle of the Ga catalyst droplet for the structural transition is deduced.

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.

Responses of photosynthesis and component processes to drought and temperature stress: are Mediterranean trees fit for climate change?

Global warming is raising concerns about the acclimatory capacity of trees and forests, especially in Mediterranean-type ecosystems. The sensitivity of photosynthesis to temperature is a key uncertainty for projecting the magnitude of terrestrial feedbacks on future climate change. While boreal, temperate and tropical species have been comparatively well investigated, our study provides the first comprehensive overview of the seasonal acclimatory responses of photosynthesis and its component processes to temperature in four Mediterranean climax species under natural conditions. We quantified seasonal changes in the responses of net photosynthesis (Anet), stomatal conductance (gs), mesophyllic conductance (gm) and electron-transport rate (Jcf), and investigated their sensitivity to drought and temperature stress in sunlit and shaded leaves of four Mediterranean tree species (Quercus ilex L., Pinus halepensis Mill., Arbutus unedo L. and Quercus pubescens Willd.). Sunlit leaves, but not shaded leaves, showed a pronounced seasonality in the temperature responses of Anet, gs, gm and Jcf. All four species and variables showed a remarkably dynamic and consistent acclimation of the thermal optimum (Topt), reaching peaks in summer ~29-32 °C. Changes in the shape of the response curves were, however, highly species-specific. Under severe drought, Topt of all variables were on average 22-29% lower. This was accompanied by narrower response curves above all in P. halepensis, reducing the optimal range for photosynthesis to the cooler morning or evening periods. Wider temperature-response curves and less strict stomatal control under severe drought were accompanied by wilting and drought-induced leaf shedding in Q. ilex and Q. pubescens and by additional branch dieback in A. unedo. Mild winter conditions led to a high Topt (~19.1-22.2 °C), benefitting the evergreen species, especially P. halepensis. Seasonal acclimation of Anet was explained better by gs and gm being less pronounced in Jcf. Drought was thus a key factor, in addition to growth temperature, to explain seasonal acclimation of photosynthesis. Severe drought periods may exceed more frequently the high acclimatory capacity of Mediterranean trees to high ambient temperatures, which could lead to reduced growth, increased leaf shedding and, for some species such as A. unedo, increased mortality risk.

Growth optimization and characterization of regular arrays of GaAs/AlGaAs core/shell nanowires for tandem solar cells on silicon.

With a band gap value of 1.7 eV, Al0.2Ga0.8As is one of the ideal III-V alloys for the development of nanowire-based Tandem Solar Cells on silicon. Nevertheless, growing self-catalysed AlGaAs nanowires on silicon by solid-source molecular beam epitaxy is a very difficult task due to the oxidation of Al adatoms by the SiO2 layer present on the surface. Here we propose a nanowire structure including a p.i.n radial junction inside an Al0.2Ga0.8As shell grown on a p-GaAs core. The crystalline structure of such self-catalysed nanowires grown on an epi-ready Si(111) substrate (with a thin native SiO2 layer) was investigated by transmission electronic microscopy and photoluminescence. I(V) measurements performed on single nanowires have shown a diode-like behaviour corresponding to the radial p.i.n junction inside the Al0.2Ga0.8As shell. Moreover, a current generation under the electron beam was evidenced over the entire radial junction along the nanowires by means of electron beam induced current (EBIC) microscopy. The same structure was reproduced on patterned substrates with a SiO2 mask, producing an ordered hexagonal array. High and uniform yields from 83% to 87% of vertical nanowires were obtained on 0.9 × 0.9 cm2 patterned areas. EBIC mapping performed on these nanowires confirmed the good electrical properties of the radial junction within the nanowires.

Author Correction: Crowther et al. reply.

In this Brief Communications Arising Reply, the affiliation for author P. H. Templer was incorrectly listed as 'Department of Ecology & Evolutionary Biology, University of California Irvine, Irvine, California 92697, USA' instead of 'Department of Biology, Boston University, Boston, Massachusetts 02215, USA'. This has been corrected online.

Publisher Correction: Large anisotropy of ferroelectric and pyroelectric properties in heteroepitaxial oxide layers.

A correction to this article has been published and is linked from the HTML and PDF versions of this paper. The error has been fixed in the paper.

Large anisotropy of ferroelectric and pyroelectric properties in heteroepitaxial oxide layers.

Epitaxial PbZr0.52Ti0.48O3 (PZT) layers were integrated on Si(001) with single PZT {001} orientation, mosaïcity below 1° and a majority of a-oriented ferroelectric domains (∼65%). Ferroelectric and pyroelectric properties are determined along both the out-of-plane and in-plane directions through parallel-plate capacitor and coplanar interdigital capacitor along the <100>PZT direction. A large anisotropy in these properties is observed. The in-plane remnant polarization (21.5 µC.cm-2) is almost twice larger than that measured along the out-of-plane direction (13.5 µC.cm-2), in agreement with the domain orientation. Oppositely, the in-plane pyroelectric coefficient (-285 µC.m-2.K-1) is much lower than that measured out-of-plane (-480 µC.m-2.K-1). The pyroelectric anisotropy is explicated in term of degree of structural freedom with temperature. In particular, the low in-plane pyroelectric coefficient is explained by a two-dimensional clamping of the layers on the substrate which induces tensile stress (from thermal expansion), competing with the decreasing tetragonality of a-domains (shortening of the polar c-axis lattice parameter). Temperature-dependent XRD measurements have revealed an increased fraction of a-domains with temperature, attesting the occurrence of a partial two-dimensional clamping. These observed properties are of critical importance for integrated pyroelectric devices.

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.

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.

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.

Quantifying global soil carbon losses in response to warming.

The majority of the Earth's terrestrial carbon is stored in the soil. If anthropogenic warming stimulates the loss of this carbon to the atmosphere, it could drive further planetary warming. Despite evidence that warming enhances carbon fluxes to and from the soil, the net global balance between these responses remains uncertain. Here we present a comprehensive analysis of warming-induced changes in soil carbon stocks by assembling data from 49 field experiments located across North America, Europe and Asia. We find that the effects of warming are contingent on the size of the initial soil carbon stock, with considerable losses occurring in high-latitude areas. By extrapolating this empirical relationship to the global scale, we provide estimates of soil carbon sensitivity to warming that may help to constrain Earth system model projections. Our empirical relationship suggests that global soil carbon stocks in the upper soil horizons will fall by 30 ± 30 petagrams of carbon to 203 ± 161 petagrams of carbon under one degree of warming, depending on the rate at which the effects of warming are realized. Under the conservative assumption that the response of soil carbon to warming occurs within a year, a business-as-usual climate scenario would drive the loss of 55 ± 50 petagrams of carbon from the upper soil horizons by 2050. This value is around 12-17 per cent of the expected anthropogenic emissions over this period. Despite the considerable uncertainty in our estimates, the direction of the global soil carbon response is consistent across all scenarios. This provides strong empirical support for the idea that rising temperatures will stimulate the net loss of soil carbon to the atmosphere, driving a positive land carbon-climate feedback that could accelerate climate change.

Relatively stable response of fruiting stage to warming and cooling relative to other phenological events.

The timing of the fruit-set stage (i.e., start and end of fruit set) is crucial in a plant's life cycle, but its response to temperature change is still unclear. We investigated the timing of seven phenological events, including fruit-set dates during 3 yr for six alpine plants transplanted to warmer (approximately +3.5°C in soils) and cooler (approximately -3.5°C in soils) locations along an altitudinal gradient in the Tibetan area. We found that fruit-set dates remained relatively stable under both warming and cooling during the 3-yr transplant experiment. Three earlier phenological events (emergence of first leaf, first bud set, and first flowering) and two later phenological events (first leaf coloring and complete leaf coloring) were earlier by 4.8-8.2 d/°C and later by 3.2-7.1 d/°C in response to warming. Conversely, cooling delayed the three earlier events by 3.8-6.9 d/°C and advanced the two later events by 3.2-8.1 d/°C for all plant species. The timing of the first and/or last fruit-set dates, however, did not change significantly compared to earlier and later phenological events. Statistical analyses also showed that the dates of fruit set were not significantly correlated or had lower correlations with changes of soil temperature relative to the earlier and later phenological events. Alpine plants may thus acclimate to changes in temperature for their fruiting function by maintaining relatively stable timings of fruit set compared with other phenological events to maximize the success of seed maturation and dispersal in response to short-term warming or cooling.