(2015) The application of ecological stoichiometry to plant-microbial-soil organic matter transformations. . : -.LinkDoi: 10.1890/14-0777.1
Blanch J.S., Peñuelas J., Llusià J., Sardans J., Owen S.M. (2015) Differences in photosynthesis and terpene content in leaves and roots of wild-type and transgenic Arabidopsis thaliana plants. Russian Journal of Plant Physiology. 62: 823-829.LinkDoi: 10.1134/S1021443715060035
We investigated the hypotheses that two different varieties of Arabidopsis thaliana show differences in physiology and terpene production. The two varieties of A. thaliana used in this study were wild-type (WT) and transgenic line (CoxIV-FaNES I) genetically modified to emit nerolidol with linalool/nerolidol synthase (COX). Photosynthetic rate, electron transport rate, fluorescence, leaf volatile terpene contents and root volatile terpene contents were analyzed. For both types, we found co-eluting α-pinene+β-ocimene, limonene, and humulene in leaves; and in the roots we found co-eluting α-pinene+β-ocimene, sabinene+β-pinene, β-myrcene, limonene, and humulene. At the end of the growing cycle, COX plants tended to have lower pools of terpene compounds in their leaves, with 78.6% lower photosynthesis rates and 30.8% lower electron transport rates, compared with WT plants at that time. The maximal photochemical efficiency Fv/Fm was also significantly lower (25.5%) in COX plants, indicating that these varieties were more stressed than WT plants. However, COX plants had higher (239%) root terpene contents compared to WT plants. COX plants appear to favor root production of volatile terpenes rather than leaf production. Thus we conclude that there were significant differences between COX and WT plants in terms of terpenoid pools, stress status and physiology. © 2015, Pleiades Publishing, Ltd.
Fernández-Martínez M., Vicca S., Janssens I.A., Sardans J., Luyssaert S., Campioli M., Chapin F.S., Ciais P., Malhi Y., Obersteiner M., Papale D., Piao S.L., Reichstein M., Rodà F., Peñuelas J. (2015) Reply to 'Uncertain effects of nutrient availability on global forest carbon balance' and 'Data quality and the role of nutrients in forest carbon-use efficiency'. Nature Climate Change. 5: 960-961.LinkDoi: 10.1038/nclimate2794
[No abstract available]
Gargallo-Garriga A., Sardans J., Perez-Trujillo M., Oravec M., Urban O., Jentsch A., Kreyling J., Beierkuhnlein C., Parella T., Penuelas J. (2015) Warming differentially influences the effects of drought on stoichiometry and metabolomics in shoots and roots. New Phytologist. : 0-0.LinkDoi: 10.1111/nph.13377
Plants in natural environments are increasingly being subjected to a combination of abiotic stresses, such as drought and warming, in many regions. The effects of each stress and the combination of stresses on the functioning of shoots and roots have been studied extensively, but little is known about the simultaneous metabolome responses of the different organs of the plant to different stresses acting at once. We studied the shift in metabolism and elemental composition of shoots and roots of two perennial grasses, Holcus lanatus and Alopecurus pratensis, in response to simultaneous drought and warming. These species responded differently to individual and simultaneous stresses. These responses were even opposite in roots and shoots. In plants exposed to simultaneous drought and warming, terpenes, catechin and indole acetic acid accumulated in shoots, whereas amino acids, quinic acid, nitrogenous bases, the osmoprotectants choline and glycine betaine, and elements involved in growth (nitrogen, phosphorus and potassium) accumulated in roots. Under drought, warming further increased the allocation of primary metabolic activity to roots and changed the composition of secondary metabolites in shoots. These results highlight the plasticity of plant metabolomes and stoichiometry, and the different complementary responses of shoots and roots to complex environmental conditions. © 2015 New Phytologist Trust.
Huang Z., Liu B., Davis M., Sardans J., Peñuelas J., Billings S. (2015) Long-term nitrogen deposition linked to reduced water use efficiency in forests with low phosphorus availability. New Phytologist. : 0-0.LinkDoi: 10.1111/nph.13785
The impact of long-term nitrogen (N) deposition is under-studied in phosphorus (P)-limited subtropical forests. We exploited historically collected herbarium specimens to investigate potential physiological responses of trees in three subtropical forests representing an urban-to-rural gradient, across which N deposition has probably varied over the past six decades. We measured foliar [N] and [P] and stable carbon (δ13C), oxygen (δ18O) and nitrogen (δ15N) isotopic compositions in tissue from herbarium specimens of plant species collected from 1947 to 2014. Foliar [N] and N: P increased, and δ15N and [P] decreased in the two forests close to urban centers. Consistent with recent studies demonstrating that N deposition in the region is 15N-depleted, these data suggest that the increased foliar [N] and N: P, and decreased [P], may be attributable to atmospheric deposition and associated enhancement of P limitation. Estimates of intrinsic water use efficiency calculated from foliar δ13C decreased by c. 30% from the 1950s to 2014, contrasting with multiple studies investigating similar parameters in N-limited forests. This effect may reflect decreased photosynthesis, as suggested by a conceptual model of foliar δ13C and δ18O. Long-term N deposition may exacerbate P limitation and mitigate projected increases in carbon stocks driven by elevated CO2 in forests on P-limited soils. © 2015 New Phytologist Trust.
Rajsnerova P., Klem K., Holub P., Novotna K., Vecerova K., Kozacikova M., Rivas-Ubach A., Sardans J., Marek M.V., Penuelas J., Urban O. (2015) Morphological, biochemical and physiological traits of upper and lower canopy leaves of European beech tend to converge with increasing altitude. Tree Physiology. 35: 47-60.LinkDoi: 10.1093/treephys/tpu104
The present work has explored for the first time acclimation of upper versus lower canopy leaves along an altitudinal gradient. We tested the hypothesis that restrictive climatic conditions associated with high altitudes reduce within-canopy variations of leaf traits. The investigated beech (Fagus sylvatica L.) forest is located on the southern slope of the Hrubý Jeseník Mountains (Czech Republic). All measurements were taken on leaves from upper and lower parts of the canopy of mature trees (>85 years old) growing at low (400 m above sea level, a.s.l.), middle (720 m a.s.l.) and high (1100 m a.s.l.) altitudes. Compared with trees at higher altitudes, those growing at low altitudes had lower stomatal conductance, slightly lower CO2 assimilation rate (A max) and leaf mass per area (LMA), and higher photochemical reflectance index, water-use efficiency and Rubisco content. Given similar stand densities at all altitudes, the different growth conditions result in a more open canopy and higher penetration of light into lower canopy with increasing altitude. Even though strong vertical gradients in light intensity occurred across the canopy at all altitudes, lower canopy leaves at high altitudes tended to acquire the same morphological, biochemical and physiological traits as did upper leaves. While elevation had no significant effect on nitrogen (N) and carbon (C) contents per unit leaf area, LMA, or total content of chlorophylls and epidermal flavonoids in upper leaves, these increased significantly in lower leaves at higher altitudes. The increases in N content of lower leaves were coupled with similar changes in A max. Moreover, a high N content coincided with high Rubisco concentrations in lower but not in upper canopy leaves. Our results show that the limiting role of light in lower parts of the canopy is reduced at high altitudes. A great capacity of trees to adjust the entire canopy is thus demonstrated. © © The Author 2015. Published by Oxford University Press. All rights reserved.
Rivas-Ubach A., Sardans J., Hódar J.A., Garcia-Porta J., Guenther A., Oravec M., Urban O., Peñuelas J. (2015) Similar local, but different systemic, metabolomic responses of closely related pine subspecies to folivory by caterpillars of the processionary moth. Plant Biology. : 0-0.LinkDoi: 10.1111/plb.12422
Plants respond locally and systemically to herbivore attack. Most of the research conducted on plant-herbivore relationships at element and molecular levels have focused on the elemental composition or/and certain molecular compounds or specific families of defence metabolites showing that herbivores tend to select plant individuals or species with higher nutrient concentrations and avoid those with higher levels of defence compounds. We performed stoichiometric and metabolomics, both local and systemic, analyses in two subspecies of Pinus sylvestris under attack from caterpillars of the pine processionary moth, an important pest in the Mediterranean Basin. Both pine subspecies responded locally to folivory mainly by increasing relative concentrations of terpenes and some phenolics. Systemic responses differed between pine subspecies, and most of the metabolites presented intermediate concentrations between those of the affected parts and unattacked trees. Our results support the hypothesis that foliar nutrient concentrations are not a key factor for plant selection by adult female processionary moths for oviposition, since folivory was not associated with any of the elements analysed. Phenolic compounds generally did not increase in the attacked trees, questioning the suggestion of induction of phenolics following folivory attack and the anti-feeding properties of phenolics. Herbivory attack produced a general systemic shift in pines, in both primary and secondary metabolism, which was less intense and chemically different from the local responses. Local pine responses were similar between pine subspecies, while systemic responses were more distant. © 2015 German Botanical Society and The Royal Botanical Society of the Netherlands.
Sardans J., Alonso R., Janssens I.A., Carnicer J., Vereseglou S., Rillig M.C., Fernández-Martínez M., Sanders T.G.M., Peñuelas J. (2015) Foliar and soil concentrations and stoichiometry of nitrogen and phosphorous across European Pinus sylvestris forests: Relationships with climate, N deposition and tree growth. Functional Ecology. : 0-0.LinkDoi: 10.1111/1365-2435.12541
This study investigated the factors underlying the variability of needle and soil elemental composition and stoichiometry and their relationships with growth in Pinus sylvestris forests throughout the species' distribution in Europe by analysing data from 2245 forest stands. Needle N concentrations and N:P ratios were positively correlated with total atmospheric N deposition, whereas needle P concentrations were negatively correlated. These relationships were especially pronounced at sites where high levels of N deposition coincided with both higher mean annual temperature and higher mean annual precipitation. Trends towards foliar P deficiency were thus more marked when high N deposition coincided with climatic conditions favourable to plant production. Atmospheric N deposition was positively correlated with soil solution NO3- , SO42- , K+, P and Ca2+ concentrations, the soil solution NO3-:P ratio, total soil N and the total soil N:Olsen P ratio and was negatively correlated with soil Olsen P concentration. Despite these nutrient imbalances, during the period studied (1990-2006), N deposition was positively related with Pinus sylvestris absolute basal diameter (BD) growth, although only accounting for the 10% of the total variance. However, neither N deposition nor needle N concentration was related with relative annual BD growth. In contrast, needle P concentration was positively related with both absolute and relative annual BD growth. These results thus indicate a tendency of European P. sylvestris forests to store N in trees and soil in response to N deposition and unveil a trend towards increased nutrient losses in run-off as a consequence of higher soil solution N concentrations. Overall, the data show increasing ecosystem nutrient imbalances with increasingly limiting roles of P and other nutrients such as K in European P. sylvestris forests, especially in the centre of their distribution where higher levels of N deposition are observed. Thus, although the data show that N deposition has had an overall positive effect on P. sylvestris growth, the effect of continuous N deposition, associated with decreasing P and K and increasing N:P in leaves and in soil, may in the future become detrimental for the growth and competitive ability of P. sylvestris trees. © 2015 British Ecological Society.
Sardans J., Janssens I.A., Alonso R., Veresoglou S.D., Rillig M.C., Sanders T.G.M., Carnicer J., Filella I., Farre-Armengol G., Penuelas J. (2015) Foliar elemental composition of European forest tree species associated with evolutionary traits and present environmental and competitive conditions. Global Ecology and Biogeography. 24: 240-255.LinkDoi: 10.1111/geb.12253
Aim: Plant elemental composition and stoichiometry are crucial for plant structure and function. We studied to what extent elemental stoichiometry in plants might be strongly related to environmental drivers and competition from coexisting species. Location: Europe. Methods: We analysed foliar N, P, K, Ca and Mg concentrations and their ratios among 50 species of European forest trees sampled in 5284 plots across Europe and their relationships with phylogeny, forest type, current climate and N deposition. Results: Phylogeny is strongly related to overall foliar elemental composition in European tree species. Species identity explained 56.7% of the overall foliar elemental composition and stoichiometry. Forest type and current climatic conditions also partially explained the differences in foliar elemental composition among species. In the same genus co-occurring species had overall higher differences in foliar elemental composition and stoichiometry than the non-co-occurring species. Main conclusions: The different foliar elemental compositions among species are related to phylogenetic distances, but they are also related to current climatic conditions, forest types, drivers of global change such as atmospheric N deposition, and to differences among co-occurring species as a probable consequence of niche specialization to reduce direct competition for the same resources. Different species have their own 'fixed' foliar elemental compositions but retain some degree of plasticity to the current climatic and competitive conditions. A wider set of elements beyond N and P better represent the biogeochemical niche and are highly sensitive to plant function. Foliar elemental composition can thus be useful for representing important aspects of plant species niches. © 2014 John Wiley & Sons Ltd.
Sardans J., Penuelas J. (2015) Trees increase their P: N ratio with size. Global Ecology and Biogeography. 24: 147-156.LinkDoi: 10.1111/geb.12231
Aim: Phosphorus (P) tends to become limiting in ageing terrestrial ecosystems, and its resorption efficiency is higher than for other elements such as nitrogen (N). We thus hypothesized that trees should store more P than those other elements such as N when tree size increases and that this process should be enhanced in slow-growing late-successional trees. Location: Catalan forests. Methods: We used data from the Catalan Forest Inventory that contains field data on the P and N contents of total aboveground, foliar and woody biomasses of the diverse mediterranean, temperate and alpine forests of Catalonia (1018 sites). We used correlation and general linear models to analyse the allometric relationships between the nutrient contents of different aboveground biomass fractions (foliar, branches and stems) and total aboveground biomass. Results: Aboveground forest P content increases proportionally more than aboveground forest N content with increasing aboveground biomass. Two mechanisms underlie this. First, woody biomass increases proportionally more than foliar biomass, with woody biomass having a higher P:N ratio than foliar biomass. Second, the P:N ratio of wood increases with tree size. These results are consistent with the generally higher foliar resorption of P than of N. Slow-growing species accumulate more total P aboveground with size than fast-growing species, mainly as a result of their large capacity to store P in wood. Main conclusions: Trees may have developed long-term adaptive mechanisms to store P in biomass, mainly in wood, thereby slowing the loss of P from ecosystems, reducing its availability for competitors and implying an increase in the P:N ratio in forest biomass with ageing. This trend to accumulate more P than N with size is more accentuated in slow-growing, large, long-lived species of late successional stages. In this way they partly counterbalance the gradual decrease of P in the soil. © 2014 John Wiley & Sons Ltd.
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