Achotegui-Castells A., Llusia J., Hodar J.A., Penuelas J. (2013) Needle terpene concentrations and emissions of two coexisting subspecies of Scots pine attacked by the pine processionary moth (Thaumetopoea pityocampa). Acta Physiologiae Plantarum. 35: 3047-3058.LinkDoi: 10.1007/s11738-013-1337-3
Mediterranean pine forests are often attacked by caterpillars of Thaumetopoea pityocampa (Lep., Thaumetopoidae), one of the most important defoliators in the Mediterranean region causing large economic losses and ecological effects. The needle terpene concentrations and emissions may play a key role in the defense of pines. We studied two subspecies of Pinus sylvestris, nevadensis (an endemic and relict subspecies) and iberica, with different levels of caterpillar attack in Sierra Nevada mountains (Spain). GC-MS analyses showed large total concentrations of terpenes (6 to 39 mg g-1 of dry weight) in the needles of both subspecies under field conditions. Concentrations were 25 % higher in "Non-Attacked Trees" (NATs) of the iberica than in the nevadensis subspecies. The branches of NATs had terpene concentrations 20 % higher than those of "Attacked Branches of attacked trees" (ABs). Within attacked trees, the "Non-Attacked Branches" (NABs) also had terpene concentrations 20 % higher than those of ABs. Mainly α-pinene and germacrene D had higher concentrations in NATs and NABs than in ABs. Some terpenes had higher concentrations in NABs than in NATs, indicating possible systemic reactions. In subsp. nevadensis, the percentage of monoterpenes relative to total terpenes was higher in ABs than in other attack states. The rates of emission in nevadensis (standardized to 30 °C) were ca. three times higher in ABs than in NABs and NATs. These results suggest that the lower terpene concentrations and high percentages of monoterpenes in ABs were produced by a combination of emission losses and terpene induction in response to herbivorous attack. © 2013 Franciszek Górski Institute of Plant Physiology, Polish Academy of Sciences, Kraków.
Farré-Armengol G., Filella I., Llusia J., Peñuelas J. (2013) Floral volatile organic compounds: Between attraction and deterrence of visitors under global change. Perspectives in Plant Ecology, Evolution and Systematics. 15: 56-67.LinkDoi: 10.1016/j.ppees.2012.12.002
Plants produce and emit a large variety of volatile organic compounds that play key roles in interactions with abiotic and biotic environments. One of these roles is the attraction of animals (mainly insects) that act as vectors of pollen to ensure reproduction. Here we update the current knowledge of four key aspects of floral emissions: (1) the relative importance and interaction of olfactory signals and visual cues, (2) the spatial and temporal patterns of emission in flowers, (3) the attractive and defensive functions of floral volatiles and their interference, and (4) the effects of global change on floral emissions and plant-pollinator interactions. Finally, we propose future lines of research in this field that need to be addressed or investigated further. © 2012 Perspectives in Plant Ecology, Evolution and Systematics.
Filella I., Primante C., Llusia J., Martin Gonzalez A.M., Seco R., Farre-Armengol G., Rodrigo A., Bosch J., Penuelas J. (2013) Floral advertisement scent in a changing plant-pollinators market. Scientific Reports. 3: 0-0.LinkDoi: 10.1038/srep03434
Plant-pollinator systems may be considered as biological markets in which pollinators choose between different flowers that advertise their nectar/pollen rewards. Although expected to play a major role in structuring plant-pollinator interactions, community-wide patterns of flower scent signals remain largely unexplored. Here we show for the first time that scent advertisement is higher in plant species that bloom early in the flowering period when pollinators are scarce relative to flowers than in species blooming later in the season when there is a surplus of pollinators relative to flowers. We also show that less abundant flowering species that may compete with dominant species for pollinator visitation early in the flowering period emit much higher proportions of the generalist attractant β-ocimene. Overall, we provide a first community-wide description of the key role of seasonal dynamics of plant-specific flower scent emissions, and reveal the coexistence of contrasting plant signaling strategies in a plant-pollinator market.
Llusia J., Penuelas J., Guenther A., Rapparini F. (2013) Seasonal variations in terpene emission factors of dominant species in four ecosystems in NE Spain. Atmospheric Environment. 70: 149-158.LinkDoi: 10.1016/j.atmosenv.2013.01.005
We studied the daily patterns in the rates of foliar terpene emissions by four typical species from the Mediterranean region in two days of early spring and two days of summer in 4 localities of increasing biomass cover in Northern Spain. The species studied were Thymelaea tinctoria (in Monegros), Quercus coccifera (in Garraf), Quercus ilex (in Prades) and Fagus sylvatica (in Montseny). Of the total 43 VOCs detected, 23 were monoterpenes, 5 sesquiterpenes and 15 were not terpenes. Sesquiterpenes were the main terpenes emitted from T. tinctoria. Total VOC emission rates were on average about 15 times higher in summer than in early spring. The maximum rates of emission were recorded around midday. Emissions nearly stopped in the dark. No significant differences were found for nocturnal total terpene emission rates between places and seasons. The seasonal variations in the rate of terpene emissions and in their chemical composition can be explained mainly by dramatic changes in emission factors (emission capacity) associated in some cases, such as for beech trees, with very different foliar ontogenical characteristics between spring and summer. The results show that temperature and light-standardised emission rates were on average about 15 times higher in summer than in early spring, which, corroborating other works, calls to attention when applying the same emission factor in modelling throughout the different seasons of the year. © 2013 Elsevier Ltd.
Penuelas J., Guenther A., Rapparini F., Llusia J., Filella I., Seco R., Estiarte M., Mejia-Chang M., Ogaya R., Ibanez J., Sardans J., Castano L.M., Turnipseed A., Duhl T., Harley P., Vila J., Estavillo J.M., Menendez S., Facini O., Baraldi R., Geron C., Mak J., Patton E.G., Jiang X., Greenberg J. (2013) Intensive measurements of gas, water, and energy exchange between vegetation and troposphere during the MONTES campaign in a vegetation gradient from short semi-desertic shrublands to tall wet temperate forests in the NW Mediterranean Basin. Atmospheric Environment. 75: 348-364.LinkDoi: 10.1016/j.atmosenv.2013.04.032
MONTES ("Woodlands") was a multidisciplinary international field campaign aimed at measuring energy, water and especially gas exchange between vegetation and atmosphere in a gradient from short semi-desertic shrublands to tall wet temperate forests in NE Spain in the North Western Mediterranean Basin (WMB). The measurements were performed at a semidesertic area (Monegros), at a coastal Mediterranean shrubland area (Garraf), at a typical Mediterranean holm oak forest area (Prades) and at a wet temperate beech forest (Montseny) during spring (April 2010) under optimal plant physiological conditions in driest-warmest sites and during summer (July 2010) with drought and heat stresses in the driest-warmest sites and optimal conditions in the wettest-coolest site. The objective of this campaign was to study the differences in gas, water and energy exchange occurring at different vegetation coverages and biomasses. Particular attention was devoted to quantitatively understand the exchange of biogenic volatile organic compounds (BVOCs) because of their biological and environmental effects in the WMB. A wide range of instruments (GC-MS, PTR-MS, meteorological sensors, O3 monitors,. .) and vertical platforms such as masts, tethered balloons and aircraft were used to characterize the gas, water and energy exchange at increasing footprint areas by measuring vertical profiles. In this paper we provide an overview of the MONTES campaign: the objectives, the characterization of the biomass and gas, water and energy exchange in the 4 sites-areas using satellite data, the estimation of isoprene and monoterpene emissions using MEGAN model, the measurements performed and the first results. The isoprene and monoterpene emission rates estimated with MEGAN and emission factors measured at the foliar level for the dominant species ranged from about 0 to 0.2mgm-2h-1 in April. The warmer temperature in July resulted in higher model estimates from about 0 to ca. 1.6mgm-2h-1 for isoprene and ca. 4.5mgm-2h-1 for monoterpenes, depending on the site vegetation and footprint area considered. There were clear daily and seasonal patterns with higher emission rates and mixing ratios at midday and summer relative to early morning and early spring. There was a significant trend in CO2 fixation (from 1 to 10mgCm-2d-1), transpiration (from1-5kgCm-2d-1), and sensible and latent heat from the warmest-driest to the coolest-wettest site. The results showed the strong land-cover-specific influence on emissions of BVOCs, gas, energy and water exchange, and therefore demonstrate the potential for feed-back to atmospheric chemistry and climate. •We present a multidisciplinary biosphere-atmosphere field campaign.•We measured a gradient from semi-desertic shrublands to wet temperate forests.•A wide range of instruments and vertical platforms were used.•Land cover strongly influenced emissions of BVOCs and gas, energy and water exchange.•Vegetation has strong potential for feed-back to atmospheric chemistry and climate. © 2013 Elsevier Ltd.
Penuelas J., Poulter B., Sardans J., Ciais P., Van Der Velde M., Bopp L., Boucher O., Godderis Y., Hinsinger P., Llusia J., Nardin E., Vicca S., Obersteiner M., Janssens I.A. (2013) Human-induced nitrogen-phosphorus imbalances alter natural and managed ecosystems across the globe. Nature Communications. 4: 0-0.LinkDoi: 10.1038/ncomms3934
The availability of carbon from rising atmospheric carbon dioxide levels and of nitrogen from various human-induced inputs to ecosystems is continuously increasing; however, these increases are not paralleled by a similar increase in phosphorus inputs. The inexorable change in the stoichiometry of carbon and nitrogen relative to phosphorus has no equivalent in Earth's history. Here we report the profound and yet uncertain consequences of the human imprint on the phosphorus cycle and nitrogen:phosphorus stoichiometry for the structure, functioning and diversity of terrestrial and aquatic organisms and ecosystems. A mass balance approach is used to show that limited phosphorus and nitrogen availability are likely to jointly reduce future carbon storage by natural ecosystems during this century. Further, if phosphorus fertilizers cannot be made increasingly accessible, the crop yields projections of the Millennium Ecosystem Assessment imply an increase of the nutrient deficit in developing regions. © 2013 Macmillan Publishers Limited.
Penuelas J., Sardans J., Estiarte M., Ogaya R., Carnicer J., Coll M., Barbeta A., Rivas-Ubach A., Llusia J., Garbulsky M., Filella I., Jump A.S. (2013) Evidence of current impact of climate change on life: A walk from genes to the biosphere. Global Change Biology. 19: 2303-2338.LinkDoi: 10.1111/gcb.12143
We review the evidence of how organisms and populations are currently responding to climate change through phenotypic plasticity, genotypic evolution, changes in distribution and, in some cases, local extinction. Organisms alter their gene expression and metabolism to increase the concentrations of several antistress compounds and to change their physiology, phenology, growth and reproduction in response to climate change. Rapid adaptation and microevolution occur at the population level. Together with these phenotypic and genotypic adaptations, the movement of organisms and the turnover of populations can lead to migration toward habitats with better conditions unless hindered by barriers. Both migration and local extinction of populations have occurred. However, many unknowns for all these processes remain. The roles of phenotypic plasticity and genotypic evolution and their possible trade-offs and links with population structure warrant further research. The application of omic techniques to ecological studies will greatly favor this research. It remains poorly understood how climate change will result in asymmetrical responses of species and how it will interact with other increasing global impacts, such as N eutrophication, changes in environmental N : P ratios and species invasion, among many others. The biogeochemical and biophysical feedbacks on climate of all these changes in vegetation are also poorly understood. We here review the evidence of responses to climate change and discuss the perspectives for increasing our knowledge of the interactions between climate change and life. © 2013 John Wiley & Sons Ltd.
Penuelas J., Sardans J., Llusia J., Silva J., Owen S.M., Bala-Ola B., Linatoc A.C., Dalimin M.N., Niinemets U. (2013) Foliar chemistry and standing folivory of early and late-successional species in a Bornean rainforest. Plant Ecology and Diversity. 6: 245-256.LinkDoi: 10.1080/17550874.2013.768713
Background: Few studies have investigated the chemical, morphological and physiological foliar traits and the intensity of standing folivory in a representative set of species of tropical rainforests including species of different successional stages. Aims: (i) To quantify leaf elemental composition, leaf phenolics and tannin concentrations, physical leaf traits and the intensity of standing folivory in a representative set of species of different successional stages in a Bornean tropical rainforest, and (ii) to investigate the relationships among leaf traits and between leaf traits and accumulated standing folivory. Methods: Analyses of leaf elemental concentrations, phenolics (Ph) and tannin (Tan) concentrations, leaf mass area (LMA), C assimilation rate and accumulated standing folivory in 88 common rainforest species of Borneo. Results and Conclusions: Accumulated standing folivory was correlated with the scores of the first axis of the elemental concentrations principal component analysis (mainly loaded by K and C:K and N:K ratios) with lower accumulated standing folivory at high leaf K concentrations (R = -0.33, P = 0.0016). The results show that consistent with growth rate hypothesis, fast-growing pioneer species have lower leaf N:P ratios than late-successional species, that species with higher leaf N concentration have lower LMA according with the 'leaf economics spectrum' hypothesis, and that species with lower leaf nutrient concentration allocate more C to leaf phenolics. This study also shows that species with different ecological roles have different biogeochemical 'niches' assessed as foliar elemental composition. © 2013 Copyright 2013 Botanical Society of Scotland and Taylor & Francis.
Peñuelas J., Marino G., Llusia J., Morfopoulos C., Farré-Armengol G., Filella I. (2013) Photochemical reflectance index as an indirect estimator of foliar isoprenoid emissions at the ecosystem level. Nature Communications. 4: 0-0.LinkDoi: 10.1038/ncomms3604
Terrestrial plants re-emit around 1-2% of the carbon they fix as isoprene and monoterpenes. These emissions have major roles in the ecological relationships among living organisms and in atmospheric chemistry and climate, and yet their actual quantification at the ecosystem level in different regions is far from being resolved with available models and field measurements. Here we provide evidence that a simple remote sensing index, the photochemical reflectance index, which is indicative of light use efficiency, is a good indirect estimator of foliar isoprenoid emissions and can therefore be used to sense them remotely. These results open new perspectives for the potential use of remote sensing techniques to track isoprenoid emissions from vegetation at larger scales. On the other hand, our study shows the potential of this photochemical reflectance index technique to validate the availability of photosynthetic reducing power as a factor involved in isoprenoid production. © 2013 Macmillan Publishers Limited. All rights reserved.
Seco R., Penuelas J., Filella I., Llusia J., Schallhart S., Metzger A., Muller M., Hansel A. (2013) Volatile organic compounds in the western Mediterranean basin: Urban and rural winter measurements during the DAURE campaign. Atmospheric Chemistry and Physics. 13: 4291-4306.LinkDoi: 10.5194/acp-13-4291-2013
Atmospheric volatile organic compounds (VOCs) have key environmental and biological roles, but little is known about the daily VOC mixing ratios in Mediterranean urban and natural environments. We measured VOC mixing ratios concurrently at an urban and a rural site during the winter DAURE campaign in the northeastern Iberian Peninsula, by means of PTR-MS at both locations: a PTR-Quad-MS at the urban site and a PTR-ToF-MS at the rural site. All VOC mixing ratios measured were higher at the urban site (e.g. acetaldehyde, isoprene, benzene, and toluene with averages up to 1.68, 0.31, 0.58 and 2.71 ppbv, respectively), with the exception of some short-chain oxygenated VOCs such as acetone (with similar averages of 0.7-1.6 ppbv at both sites). The average diurnal pattern also differed between the sites. Most of the VOCs at the urban location showed their highest mixing ratios in the morning and evening. These peaks coincided with traffic during rush hour, the main origin of most of the VOCs analyzed. Between these two peaks, the sea breeze transported the urban air inland, thus helping to lower the VOC loading at the urban site. At the rural site, most of the measured VOCs were advected by the midday sea breeze, yielding the highest daily VOC mixing ratios (e.g. acetaldehyde, isoprene, benzene, and toluene with averages up to 0.65, 0.07, 0.19, and 0.41 ppbv, respectively). Only biogenic monoterpenes showed a clear local origin at this site. In addition, the concentrations of fine particulate matter observed at both sites, together with the synoptic meteorological conditions and radio-sounding data, allowed the identification of different atmospheric scenarios that had a clear influence on the measured VOC mixing ratios. These results highlight the differences and relationships in VOC mixing ratios between nearby urban and rural areas in Mediterranean regions. Further research in other urban-rural areas is warranted to better understand the urban-rural influence on atmospheric VOC mixing ratios under different atmospheric conditions. © 2013 Author(s).
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