Wang W.-Q., Wang C., Sardans J., Zeng C.-S., Tong C., Penuelas J. (2015) Plant invasive success associated with higher N-use efficiency and stoichiometric shifts in the soil–plant system in the Minjiang River tidal estuarine wetlands of China. Wetlands Ecology and Management. : 0-0.EnlaceDoi: 10.1007/s11273-015-9425-3
The tidal estuarine wetlands of China are rich in plant diversity, but several human-driven processes, such as species invasion, can affect the biogeochemical cycles of these ecosystems, and by changing soil conditions can inhibit the regeneration of native vegetation. We seasonally analyzed the carbon (C), nitrogen (N) and phosphorus (P) concentrations in soils and in leaves, stems and roots of the invasive species Spartina alterniflora and of the native species Cyperus malaccensis var. brevifolius Boeckeler. This latter species was analyzed both in natural non-invaded stands and in stands that had been invaded by Spartina but from which it had been removed and replaced by Cyperus. The aim was to investigate the effect of plant invasion, subsequent removal and replanting with a native species on C, N and P stoichiometry of the plant–soil system in the tidal wetlands of the Minjiang River. C and N concentrations averaged across seasons did not differ significantly among the plant species. P concentration was lower in the stems of Spartina than in the stems of the native species Cyperus but was not significantly different in the roots of the two species. The soil C and N concentrations were higher in the Spartina stand than in the Cyperus stand, whereas the soil P concentrations were not significantly different. The invasive species had a higher N-resorption capacity, N:P ratios in stem and roots, biomass, absolute growth and biomass N and had a lower relative growth rate and litter production than the native species. After the removal of the invasive plants, the regenerating native plants have a higher capacity to resorb N and lower relative growth rates. All these traits show that a conservative strategy and a high N-use efficiency and internal plant control of the N in the ecosystem underlie the invasive success of Spartina in this N-limited wetland. Relative growth rate was associated with lower plant N:P ratios, whereas absolute growth rate was associated with higher nutrient-use efficiency and lower C and N turnover and storage capacities in the biomass. Changes in soil properties produced by the establishment of an invasive plant can condition the later regeneration of native plants. © 2015 Springer Science+Business Media Dordrecht
Wang W.Q., Sardans J., Wang C., Zeng C.S., Tong C., Asensio D., Penuelas J. (2015) Ecological stoichiometry of C, N, and P of invasive Phragmites australis and native Cyperus malaccensis species in the Minjiang River tidal estuarine wetlands of China. Plant Ecology. : 0-0.EnlaceDoi: 10.1007/s11258-015-0469-5
Tidal estuarine wetlands of China are rich in plant diversity, but several global change drivers, such as species invasion, are currently affecting the biogeochemical cycles of these ecosystems. We seasonally analyzed the carbon (C), nitrogen (N), and phosphorus (P) concentrations in litters and soils and in leaves, stems, and roots of the C3 invasive species Phragmites australis (Cav.) Trin. ex Steud. and of the C4 native species Cyperus malaccensis var. brevifolius Boeckeler to investigate the effect of C3 plant invasion on C, N, and P stoichiometry in the C4 plant-dominated tidal wetlands of the Minjiang River. When averaged across seasons, the invasive species P. australis had higher N concentrations and lower P concentrations in leaves than the native species C. malaccensis. N and P concentrations were lower in litter (stem and leaf), whereas C concentrations in leaf litter were higher in P. australis than in C. malaccensis. The C, N, and P concentrations of the soil also did not differ, but plants had a lower C:N and much higher N:P ratios than soils. Root C:P and N:P ratios were lower in the growing season both in the invasive and the native species. The leaf C:N, C:P and N:P ratios peaked in summer. The invasive species had lower C:N ratio in leaves and roots, and higher N:P ratios in all biomass organs and litter than the native species, an effect related with the higher N-resorption capacity of the invasive species. Interspecific differences in C:N, C:P, and N:P ratios may likely reflect the differences in plant morphology, nutrient-use efficiency, and photosynthetic capacity between the C3 (P. australis) and C4 (C. malaccensis) plants. Our results generally suggested that the success of P. australis in these wetlands was related to its slow growth and higher resorption capacity of N and P. This implies a more conservative use of limited nutrients, particularly N, by P. australis, and to higher N concentration in its biomass thus potentially contributing to its invasiveness in these estuarine wetlands. © 2015 Springer Science+Business Media Dordrecht
Zechmeister-Boltenstern S., Keiblinger K.M., Mooshammer M., Peñuelas J., Richter A., Sardans J., Wanek W. (2015) The application of ecological stoichiometry to plant-microbial-soil organic matter transformations. Ecological Monographs. 85: 133-155.EnlaceDoi: 10.1890/14-0777.1
Elemental stoichiometry constitutes an inherent link between biogeochemistry and the structure and processes within food webs, and thus is at the core of ecosystem functioning. Stoichiometry allows for spanning different levels of biological organization, from cellular metabolism to ecosystem structure and nutrient cycling, and is therefore particularly useful for establishing links between different ecosystem compartments. We review elemental carbon : nitrogen : phosphorus (C:N:P) ratios in terrestrial ecosystems (from vegetation, leaf litter, woody debris, and dead roots, to soil microbes and organic matter). While the stoichiometry of the plant, litter, and soil compartments of ecosystems is well understood, heterotrophic microbial communities, which dominate the soil food web and drive nutrient cycling, have received increasing interest in recent years. This review highlights the effects of resource stoichiometry on soil microorganisms and decomposition, specifically on the structure and function of heterotrophic microbial communities and suggests several general patterns. First, latitudinal gradients of soil and litter stoichiometry are reflected in microbial community structure and function. Second, resource stoichiometry may cause changes in microbial interactions and community dynamics that lead to feedbacks in nutrient availability. Third, global change alters the C:N, C:P, and N:P ratios of primary producers, with repercussions for microbial decomposer communities and critical ecosystem services such as soil fertility. We argue that ecological stoichiometry provides a framework to analyze and predict such global change effects at various scales. © 2015 by the Ecological Society of America.
Aragon R., Sardans J., Penuelas J. (2014) Soil enzymes associated with carbon and nitrogen cycling in invaded and native secondary forests of northwestern Argentina. Plant and Soil. : 0-0.EnlaceDoi: 10.1007/s11104-014-2192-8
Background and aims Alien success has frequently been associated with changes in the concentrations of soil nutrients. We aim to investigate the effects of plant invasion on soil nutrients, potential enzyme activity and litter elemental composition and stoichiometry. Methods We compared stands of secondary forest invaded by Ligustrum lucidum and those dominated by natives, and performed litter chemical analyses on 3 native and 2 exotic tree species. Results Soils of invaded sites had 20 and 30 % increase in β-glucosidase and alkaline phosphatase activity, higher Olsen-phosphorus (P) and potassium (K) concentrations and lower nitrogen (N) concentration and N:P, N:K and ammonium:Olsen-P ratios. Invaded and non-invaded sites differed in their overall nutrient composition and enzyme activity. Natives and exotics differed in nine of the 16 litter elemental composition and stoichiometry variables analyzed. Conclusions The low N:P ratio in litter, the decrease in soil N in invaded stands and the low N concentration of exotics suggest that N is the limiting nutrient and that exotic success is related to higher N uptake and use efficiency. The higher investment in the acquisition of soil resources, higher nutrient uptake and use efficiency of limiting nutrients contribute to the success of exotics in this subtropical forest. © 2014 Springer International Publishing Switzerland.
Carnicer J., Sardans J., Stefanescu C., Ubach A., Bartons M., Asensio D., Penuelas J. (2014) Global biodiversity, stoichiometry and ecosystem function responses to human-induced C-N-P imbalances. Journal of Plant Physiology. : 0-0.EnlaceDoi: 10.1016/j.jplph.2014.07.022
Global change analyses usually consider biodiversity as a global asset that needs to be preserved. Biodiversity is frequently analysed mainly as a response variable affected by diverse environmental drivers. However, recent studies highlight that gradients of biodiversity are associated with gradual changes in the distribution of key dominant functional groups characterized by distinctive traits and stoichiometry, which in turn often define the rates of ecosystem processes and nutrient cycling. Moreover, pervasive links have been reported between biodiversity, food web structure, ecosystem function and species stoichiometry. Here we review current global stoichiometric gradients and how future distributional shifts in key functional groups may in turn influence basic ecosystem functions (production, nutrient cycling, decomposition) and therefore could exert a feedback effect on stoichiometric gradients. The C-N-P stoichiometry of most primary producers (phytoplankton, algae, plants) has been linked to functional trait continua (i.e. to major axes of phenotypic variation observed in inter-specific analyses of multiple traits). In contrast, the C-N-P stoichiometry of higher-level consumers remains less precisely quantified in many taxonomic groups. We show that significant links are observed between trait continua across trophic levels. In spite of recent advances, the future reciprocal feedbacks between key functional groups, biodiversity and ecosystem functions remain largely uncertain. The reported evidence, however, highlights the key role of stoichiometric traits and suggests the need of a progressive shift towards an ecosystemic and stoichiometric perspective in global biodiversity analyses.
Curiel Yuste J., Fernandez-Gonzalez A.J., Fernandez-Lopez M., Ogaya R., Penuelas J., Sardans J., Lloret F. (2014) Strong functional stability of soil microbial communities under semiarid Mediterranean conditions and subjected to long-term shifts in baseline precipitation. Soil Biology and Biochemistry. 69: 223-233.EnlaceDoi: 10.1016/j.soilbio.2013.10.045
We investigated the effect of soil microclimate on the structure and functioning of soil microbial communities in a Mediterranean Holm-oak forest subjected to 10 years of partial rain exclusion manipulations, simulating average drought conditions expected in Mediterranean areas for the following decades. We applied a high throughput DNA pyrosequencing technique coupled to parallel measurements of microbial respiration (RH) and temperature sensitivity of microbial respiration (Q10). Some consistent changes in the structure of bacterial communities suggest a slow process of community shifts parallel to the trend towards oligotrophy in response to long-term droughts. However, the structure of bacterial communities was mainly determined by short-term environmental fluctuations associated with sampling date (winter, spring and summer) rather than long-term (10 years) shifts in baseline precipitation. Moreover, long-term drought did not exert any chronic effect on the functioning of soil microbial communities (RH and Q10), emphasizing the functional stability of these communities to this long-term but mild shifts in water availability. We hypothesize that the particular conditions of the Mediterranean climate with strong seasonal shifts in both temperature and soil water availability but also characterized by very extreme environmental conditions during summer, was acting as a strong force in community assembling, selecting phenotypes adapted to the semiarid conditions characterizing Mediterranean ecosystems. Relations of climate with the phylogenetic structure and overall diversity of the communities as well as the distribution of the individual responses of different lineages (genera) to climate confirmed our hypotheses, evidencing communities dominated by thermotolerant and drought-tolerant phenotypes. © 2013 Elsevier Ltd.
Fernandez-Martinez M., Vicca S., Janssens I.A., Luyssaert S., Campioli M., Sardans J., Estiarte M., Penuelas J. (2014) Spatial variability and controls over biomass stocks, carbon fluxes, and resource-use efficiencies across forest ecosystems. Trees - Structure and Function. 28: 597-611.EnlaceDoi: 10.1007/s00468-013-0975-9
Key message: Stand age, water availability, and the length of the warm period are the most influencing controls of forest structure, functioning, and efficiency. We aimed to discern the distribution and controls of plant biomass, carbon fluxes, and resource-use efficiencies of forest ecosystems ranging from boreal to tropical forests. We analysed a global forest database containing estimates of stand biomass and carbon fluxes (400 and 111 sites, respectively) from which we calculated resource-use efficiencies (biomass production, carbon sequestration, light, and water-use efficiencies). We used the WorldClim climatic database and remote-sensing data derived from the Moderate Resolution Imaging Spectroradiometer to analyse climatic controls of ecosystem functioning. The influences of forest type, stand age, management, and nitrogen deposition were also explored. Tropical forests exhibited the largest gross carbon fluxes (photosynthesis and ecosystem respiration), but rather low net ecosystem production, which peaks in temperate forests. Stand age, water availability, and length of the warm period were the main factors controlling forest structure (biomass) and functionality (carbon fluxes and efficiencies). The interaction between temperature and precipitation was the main climatic driver of gross primary production and ecosystem respiration. The mean resource-use efficiency varied little among biomes. The spatial variability of biomass stocks and their distribution among ecosystem compartments were strongly correlated with the variability in carbon fluxes, and both were strongly controlled by climate (water availability, temperature) and stand characteristics (age, type of leaf). Gross primary production and ecosystem respiration were strongly correlated with mean annual temperature and precipitation only when precipitation and temperature were not limiting factors. Finally, our results suggest a global convergence in mean resource-use efficiencies. © 2013 Springer-Verlag Berlin Heidelberg.
Fernandez-Martinez M., Vicca S., Janssens I.A., Sardans J., Luyssaert S., Campioli M., Chapin Iii F.S., Ciais P., Malhi Y., Obersteiner M., Papale D., Piao S.L., Reichstein M., Roda F., Penuelas J. (2014) Nutrient availability as the key regulator of global forest carbon balance. Nature Climate Change. 4: 471-476.EnlaceDoi: 10.1038/nclimate2177
Forests strongly affect climate through the exchange of large amounts of atmospheric CO 2 (ref.). The main drivers of spatial variability in net ecosystem production (NEP) on a global scale are, however, poorly known. As increasing nutrient availability increases the production of biomass per unit of photosynthesis and reduces heterotrophic respiration in forests, we expected nutrients to determine carbon sequestration in forests. Our synthesis study of 92 forests in different climate zones revealed that nutrient availability indeed plays a crucial role in determining NEP and ecosystem carbon-use efficiency (CUEe; that is, the ratio of NEP to gross primary production (GPP)). Forests with high GPP exhibited high NEP only in nutrient-rich forests (CUEe = 33 ± 4%; mean ± s.e.m.). In nutrient-poor forests, a much larger proportion of GPP was released through ecosystem respiration, resulting in lower CUEe (6 ± 4%). Our finding that nutrient availability exerts a stronger control on NEP than on carbon input (GPP) conflicts with assumptions of nearly all global coupled carbon cycle-climate models, which assume that carbon inputs through photosynthesis drive biomass production and carbon sequestration. An improved global understanding of nutrient availability would therefore greatly improve carbon cycle modelling and should become a critical focus for future research. © 2014 Macmillan Publishers Limited.
Gargallo-Garriga A., Sardans J., Pérez-Trujillo M., Rivas-Ubach A., Oravec M., Vecerova K., Urban O., Jentsch A., Kreyling J., Beierkuhnlein C., Parella T., Peñuelas J. (2014) Opposite metabolic responses of shoots and roots to drought. Scientific Reports. 4: 0-0.EnlaceDoi: 10.1038/srep06829
Shoots and roots are autotrophic and heterotrophic organs of plants with different physiological functions. Do they have different metabolomes? Do their metabolisms respond differently to environmental changes such as drought? We used metabolomics and elemental analyses to answer these questions. First, we show that shoots and roots have different metabolomes and nutrient and elemental stoichiometries. Second, we show that the shoot metabolome is much more variable among species and seasons than is the root metabolome. Third, we show that the metabolic response of shoots to drought contrasts with that of roots; shoots decrease their growth metabolism (lower concentrations of sugars, amino acids, nucleosides, N, P, and K), and roots increase it in a mirrored response. Shoots are metabolically deactivated during drought to reduce the consumption of water and nutrients, whereas roots are metabolically activated to enhance the uptake of water and nutrients, together buffering the effects of drought, at least at the short term.
Greenberg J.P., Penuelas J., Guenther A., Seco R., Turnipseed A., Jiang X., Filella I., Estiarte M., Sardans J., Ogaya R., Llusia J., Rapparini F. (2014) A tethered-balloon PTRMS sampling approach for surveying of landscape-scale biogenic VOC fluxes. Atmospheric Measurement Techniques. 7: 2263-2271.EnlaceDoi: 10.5194/amt-7-2263-2014
Landscape-scale fluxes of biogenic gases were surveyed by deploying a 100 m Teflon tube attached to a tethered balloon as a sampling inlet for a fast-response proton-transfer-reaction mass spectrometer (PTRMS). Along with meteorological instruments deployed on the tethered balloon and a 3 m tripod and outputs from a regional weather model, these observations were used to estimate landscape-scale biogenic volatile organic compound fluxes with two micrometeorological techniques: mixed layer variance and surface layer gradients. This highly mobile sampling system was deployed at four field sites near Barcelona to estimate landscape-scale biogenic volatile organic compound (BVOC) emission factors in a relatively short period (3 weeks). The two micrometeorological techniques were compared with emissions predicted with a biogenic emission model using site-specific emission factors and land-cover characteristics for all four sites. The methods agreed within the uncertainty of the techniques in most cases, even though the locations had considerable heterogeneity in species distribution and complex terrain. Considering the wide range in reported BVOC emission factors for individual vegetation species (more than an order of magnitude), this temporally short and inexpensive flux estimation technique may be useful for constraining BVOC emission factors used as model inputs. © 2014 Author(s).
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