Effects of experimental warming and drought on biomass accumulation in a Mediterranean shrubland

Prieto P., Peñuelas J., Llusià J., Asensio D., Estiarte M. (2009) Effects of experimental warming and drought on biomass accumulation in a Mediterranean shrubland. Plant Ecology. 205: 179-191.
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Doi: 10.1007/s11258-009-9608-1

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We studied the effects of experimental warming and drought on the plant biomass of a Mediterranean shrubland. We monitored growth at plant level and biomass accumulation at stand level. The experimentation period stretched over 7 years (1999-2005) and we focused on the two dominant shrub species, Erica multiflora L. and Globularia alypum L. and the tree species Pinus halepensis L. The warming treatment increased shoot elongation in E. multiflora, and the drought treatment reduced shoot elongation in G. alypum. The elongation of P. halepensis remained unaffected under both treatments. The balance between the patterns observed in biomass accumulation for the three studied species in the drought plots (reduction in E. multiflora and P. halepensis and increase in G. alypum) resulted in a trend to reduce 33% the biomass of the drought treatment plots with respect to the untreated plots, which almost doubled their biomass from 1998 to 2005. The results also suggest that under drier conditions larger accumulation of dead biomass may occur at stand level, which combined with higher temperatures, may thus increase fire risk in the Mediterranean area. © Springer Science+Business Media B.V. 2009.

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The distribution of volatile isoprenoids in the soil horizons around Pinus halepensis trees

Asensio D., Owen S.M., Llusià J., Peñuelas J. (2008) The distribution of volatile isoprenoids in the soil horizons around Pinus halepensis trees. Soil Biology and Biochemistry. 40: 2937-2947.
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Doi: 10.1016/j.soilbio.2008.08.008

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We measured the terpene concentration in pentane and water extracts from soil horizons (litter, organic, top and low mineral) and from roots growing in top and low mineral horizons on a distance gradient from Pinus halepensis L. trees growing alone on a grassland. Terpene concentrations in pentane were higher than in water extracts, although β-caryophyllene showed relatively high solubility in water. The litter and roots were important sources of terpenes in soil. Alpha-pinene dominated in roots growing in both "top" (A1) and "low" (B) mineral horizons (123 ± 36 μg g-1 or 14 ± 5 mg m-2) and roots in low mineral horizon (270 ± 91 μg g-1 or 7 ± 2 mg m-2). Beta-caryophyllene dominated in litter (1469 ± 331 μg g-1 or 2004 ± 481 mg m-2). Terpene concentration in soil decreased with increasing distance to the trunk. This is likely to be related to changes in litter and roots type on the distance gradient from pine to grass and herbs. The relative contributions of all compounds, except α-pinene, were similar in the mineral soils and litter. This suggests that litter of P. halepensis is probably the main source of major terpene compounds. However, long-term emissions of α-pinene from P. halepensis roots might also contribute to α-pinene concentrations in rhizosphere soils. © 2008 Elsevier Ltd. All rights reserved.

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Interannual and seasonal changes in the soil exchange rates of monoterpenes and other VOCs in a Mediterranean shrubland

Asensio D., Peñuelas J., Prieto P., Estiarte M., Filella I., Llusià J. (2008) Interannual and seasonal changes in the soil exchange rates of monoterpenes and other VOCs in a Mediterranean shrubland. European Journal of Soil Science. 59: 878-891.
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Doi: 10.1111/j.1365-2389.2008.01057.x

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Information about soil VOC inventories and exchange rates in different soils is very scarce. Seasonality of soil VOC exchange rates is also largely unknown, despite the increasing interest in some soil volatile compounds, such as monoterpenes, because of their important role in soil ecology. We aimed to explore and quantify soil VOC exchange rates in a Mediterranean shrubland and their seasonality. Measurements of soil VOC exchange were taken using GC-MS and PTR-MS techniques, together with soil temperature, soil moisture and soil CO2 efflux measurements, during two annual campaigns with contrasting precipitation. Methanol, acetic acid, ethyl acetate, acetaldehyde, acetone, C3 and C4 carbonyls (such as methyl ethyl ketone), α-pinene and limonene, showed the highest emission rates. Maximum soil monoterpene emission rates were very low (0.003 nmol m-2 s -1) compared with foliar monoterpene emission rates. The emission rates of the other VOCs were also low (maximum 0.8 nmol m-2 s -1) except for methanol (1.2 nmol m-2 s-1). Maximum soil uptake rates for some VOCs, such as methanol and acetonitrile (ranging from -0.1 to -0.5 nmol m-2 s-1) were, however, comparable with foliar uptake rates. Further studies are needed to corroborate these results and the possible importance of the soil VOC sink in regional chemistry-climate models. Long-term severe drought increased soil monoterpene emission rates in this Mediterranean shrubland. The increases seem to be linked to changes in the soil's physical properties induced by low soil moisture. Unlike monoterpenes, other soil VOC emission rates decreased when soil moisture was low. The results suggest a seasonal control of soil temperature on the emission rates of monoterpenes and other VOCs. The emission rates increase with soil temperature. Positive correlations between the VOC exchange rates and the soil CO2 fluxes suggest that phenology of roots and microorganisms also controls seasonal changes in soil VOCs in this Mediterranean shrubland. © 2008 The Authors.

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On-line screening of soil VOCs exchange responses to moisture, temperature and root presence

Asensio D., Peñuelas J., Filella I., Llusià J. (2007) On-line screening of soil VOCs exchange responses to moisture, temperature and root presence. Plant and Soil. 291: 249-261.
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Doi: 10.1007/s11104-006-9190-4

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The exchanges of volatile organic compounds (VOCs) between soils and the atmosphere are poorly known. We investigated VOC exchange rates and how they were influenced by soil moisture, temperature and the presence of plant roots in a Mediterranean forest soil. We measured VOC exchange rates along a soil moisture gradient (5%-12.5%-20%-27.5% v/v) and a temperature gradient (10°C-15°C-25°C-35°C) using PTR-MS. Monoterpenes were identified with GC-MS. Soils were a sink rather than a source of VOCs in both soil moisture and temperature treatments (-2.16 ± 0.35 nmol m-2 s-1 and -4.90 ± 1.24 nmol m-2 s-1 respectively). Most compounds observed were oxygenated VOCs like alcohols, aldehydes and ketones and aromatic hydrocarbons. Other volatiles such as acetic acid and ethyl acetate were also observed. All those compounds had very low exchange rates (maximum uptake rates from -0.8 nmol m-2 s -1 to -0.6 nmol m-2 s-1 for methanol and acetic acid). Monoterpene exchange ranged only from -0.004 nmol m-2 s -1 to 0.004 nmol m-2 s-1 and limonene and α-pinene were the most abundant compounds. Increasing soil moisture resulted in higher soil sink activity possibly due to increases in microbial VOCs uptake activity. No general pattern of response was found in the temperature gradient for total VOCs. Roots decreased the emission of many compounds under increasing soil moisture and under increasing soil temperature. While our results showed that emission of some soil VOCs might be enhanced by the increases in soil temperature and that the uptake of most soil VOCs uptake might be reduced by the decreases of soil water availability, the low exchange rates measured indicated that soil-atmosphere VOC exchange in this system are unlikely to play an important role in atmospheric chemistry. © 2007 Springer Science+Business Media B.V.

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Interannual and interseasonal soil CO2 efflux and VOC exchange rates in a Mediterranean holm oak forest in response to experimental drought

Asensio D., Peñuelas J., Llusià J., Ogaya R., Filella I. (2007) Interannual and interseasonal soil CO2 efflux and VOC exchange rates in a Mediterranean holm oak forest in response to experimental drought. Soil Biology and Biochemistry. 39: 2471-2484.
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Doi: 10.1016/j.soilbio.2007.04.019

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Climate models predict drier conditions in the next decades in the Mediterranean basin. Given the importance of soil CO2 efflux in the global carbon balance and the important role of soil monoterpene and volatile organic compounds (VOCs) in soil ecology, we aimed to study the effects of the predicted drought on soil CO2, monoterpenes and other VOC exchange rates and their seasonal and interannual variations. We decreased soil water availability in a Mediterranean holm oak forest soil by means of an experimental drought system performed since 1999 to the present. Measurements of soil gas exchange were carried out with IRGA, GC and PTR-MS techniques during two annual campaigns of contrasting precipitation. Soil respiration was twice higher the wet year than the dry year (2.27±0.26 and 1.05±0.15, respectively), and varied seasonally from 3.76±0.85 μmol m-2 s-1 in spring, to 0.13±0.01 μmol m-2 s-1 in summer. These results highlight the strong interannual and interseasonal variation in CO2 efflux in Mediterranean ecosystems. The drought treatment produced a significant soil respiration reduction in drought plots in the wet sampling period. This reduction was even higher in wet springs (43% average reduction). These results show (1) that soil moisture is the main factor driving seasonal and interannual variations in soil respiration and (2) that the response of soil respiration to increased temperature is constrained by soil moisture. The results also show an additional control of soil CO2 efflux by physiology and phenology of trees and animals. Soil monoterpene exchange rates ranged from -0.01 to 0.004 nmol m-2 s-1, thus the contribution of this Mediterranean holm oak forest soil to the total monoterpenes atmospheric budget seems to be very low. Responses of individual monoterpenes and VOCs to the drought treatment were different depending on the compound. This suggests that the effect of soil moisture reduction in the monoterpenes and VOC exchange rates seems to be dependent on monoterpene and VOC type. In general, soil monoterpene and other VOC exchange rates were not correlated with soil CO2 efflux. In all cases, only a low proportion of variance was explained by the soil moisture changes, since almost all VOCs increased their emission rates in summer 2005, probably due to the effect of high soil temperature. Results indicate thus that physical and biological processes in soil are controlling soil VOC exchange but further research is needed on how these factors interact to produce the observed VOCs exchange responses. © 2007 Elsevier Ltd. All rights reserved.

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Seasonal soil VOC exchange rates in a Mediterranean holm oak forest and their responses to drought conditions

Asensio D., Peñuelas J., Ogaya R., Llusià J. (2007) Seasonal soil VOC exchange rates in a Mediterranean holm oak forest and their responses to drought conditions. Atmospheric Environment. 41: 2456-2466.
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Doi: 10.1016/j.atmosenv.2006.05.007

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Available information on soil volatile organic compound (VOC) exchange, emissions and uptake, is very scarce. We here describe the amounts and seasonality of soil VOC exchange during a year in a natural Mediterranean holm oak forest growing in Southern Catalonia. We investigated changes in soil VOC dynamics in drought conditions by decreasing the soil moisture to 30% of ambient conditions by artificially excluding rainfall and water runoff, and predicted the response of VOC exchange to the drought forecasted in the Mediterranean region for the next decades by GCM and ecophysiological models. The annual average of the total (detected) soil VOC and total monoterpene exchange rates were 3.2±3.2 and -0.4±0.3 μg m-2 h-1, respectively, in control plots. These values represent 0.003% of the total C emitted by soil at the study site as CO2 whereas the annual mean of soil monoterpene exchange represents 0.0004% of total C. Total soil VOC exchange rates in control plots showed seasonal variations following changes in soil moisture and phenology. Maximum values were found in spring (17±8 μg m-2 h-1). Although there was no significant global effect of drought treatment on the total soil VOC exchange rates, annual average of total VOC exchange rates in drought plots resulted in an uptake rate (-0.5±1.8 μg m-2 h-1) instead of positive net emission rates. Larger soil VOC and monoterpene exchanges were measured in drought plots than in control plots in summer, which might be mostly attributable to autotrophic (roots) metabolism. The results show that the diversity and magnitude of monoterpene and VOC soil emissions are low compared with plant emissions, that they are driven by soil moisture, that they represent a very small part of the soil-released carbon and that they may be strongly reduced or even reversed into net uptakes by the predicted decreases of soil water availability in the next decades. In all cases, it seems that VOC fluxes in soil might have greater impact on soil ecology than on atmospheric chemistry. © 2006 Elsevier Ltd. All rights reserved.

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Airborne limonene confers limited thermotolerance to Quercus ilex

Llusià J., Peñuelas J., Asensio D., Munné-Bosch S. (2005) Airborne limonene confers limited thermotolerance to Quercus ilex. Physiologia Plantarum. 123: 40-48.
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Doi: 10.1111/j.1399-3054.2004.00426.x

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The purpose of the study was to test the possible and controversial thermotolerance role of monoterpene production and emission and the related responses of antioxidants. Quercus ilex seedlings were exposed to a ramp of temperatures of 5°C steps from 25 to 50°C growing with and without limonene fumigation (7.5 μll-1). Net photosynthetic rates, maximal photochemical efficiency of PSII (Fv/Fm), oxidation state of ascorbic acid, and lipid peroxidation estimated by malondialdehyde concentrations of limonene-fumigated (LF) plants did not significantly differ from control (C) plants. No consistent changes in emissions of the other monoterpenes, α-pinene, β-phellandrene, β-pinene or β-myrcene were found. However, slight differences were found in the concentration of antioxidants. The amounts of α-tocopherol did not change or even tended to decrease at high temperatures in LF plants whereas they tended to increase by approximately 60% at 45 and 50°C relative to 25°C in C plants. Ascorbic acid reached its maximum concentration only at 45°C in LF plants whereas it reached its maximum at 35°C in C plants. β-Carotene did not decrease at high temperatures in LF plants whereas it decreased by approximately 15% at 45-50°C in C plants. Brown pigment index (BPI), an optical indicator of tissue oxidative processes, was lower in LF plants than in C plants. The photochemical reflectance index (PRI), an optical indicator of photosynthetic light use efficiency, was higher for LF plants than for C plants at elevated temperatures. Visual leaf damage (browning) tended to be less in LF plants than in C plans although not significantly (26.5 ± 8.5 versus 16.2 ± 4.8%). These results show that limonene does not confer clear and strong thermotolerance but might have some minor role. These results are in agreement with previous indications of weaker thermotolerance effect of monoterpenes than of isoprene.

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Linking isoprene with plant thermotolerance, antioxidants and monoterpene emissions

Peñuelas J., Llusià J., Asensio D., Munné-Bosch S. (2005) Linking isoprene with plant thermotolerance, antioxidants and monoterpene emissions. Plant, Cell and Environment. 28: 278-286.
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Doi: 10.1111/j.1365-3040.2004.01250.x

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The purpose of the present study was to test the possible plant thermotolerance role of isoprene and to study its relationship with non-enzymatic antioxidants and terpene emissions. The gas exchange, chlorophyll fluorescence, extent of photo- and oxidative stress, leaf damage, mechanisms of photo- and antioxidant protection, and terpene emission were measured in leaves of Quercus ilex seedlings exposed to a ramp of temperatures of 5°C steps from 25 to 50°C growing with and without isoprene (10 μL L-1) fumigation. The results showed that isoprene actually conferred thermotolerance (shifted the decrease of net photosynthetic rates from 35 to 45°C, increased Fv/Fm at 50°C from 0.38 to 0.65, and decreased the leaf area damaged from 27 to 15%), that it precluded or delayed the enhancement of the antioxidant non-enzymatic defence conferred by α-tocopherol, ascorbic acid or β-carotene consumption in response to increasing temperatures, and that it decreased by approximately 70% the emissions of monoterpenes at the highest temperatures. This suggests that there are inducible mechanisms triggered by the initial stages of thermal damage that up-regulate these antioxidant compounds at high temperatures and that these mechanisms are somehow suppressed in the presence of exogenous isoprene, which seems to already exert an antioxidant-like behaviour. © 2004 Blackwell Publishing Ltd.

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Airborne ethylene may alter antioxidant protection and reduce tolerance of holm oak to heat and drought stress

Munné-Bosch S., Peñuelas J., Asensio D., Llusià J. (2004) Airborne ethylene may alter antioxidant protection and reduce tolerance of holm oak to heat and drought stress. Plant Physiology. 136: 2937-2947.
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Doi: 10.1104/pp.104.050005

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Plant-emitted ethylene has received considerable attention as a stress hormone and is considered to play a major role at low concentrations in the tolerance of several species to biotic and abiotic stresses. However, airborne ethylene at high concentrations, such as those found in polluted areas (20-100 nL L -1) for several days, has received far less attention in studies of plant stress tolerance, though it has been shown to alter photosynthesis and reproductive stages (seed germination, flowering, and fruit ripening) in some species. To assess the potential effects of airborne ethylene on plant stress tolerance in polluted areas, the extent of oxidative stress, photo- and antioxidant protection, and visual leaf area damage were evaluated in ethylene-treated (approximately 100 nL L -1 in air) and control (without ethylene fumigation) holm oak (Quercus ilex) plants exposed to heat stress or to a combination of heat and drought stress. Control plants displayed tolerance to temperatures as high as 50°C, which might be attributed, at least in part, to enhanced xanthophyll de-epoxidation and 2-fold increases in α-tocopherol, and they suffered oxidative stress only when water deficit was superimposed on temperatures above 45°C. By contrast, ethylene-treated plants showed symptoms of oxidative stress at lower temperatures (35°C) than the controls in drought, as indicated by enhanced malondialdehyde levels, lower α-tocopherol and ascorbate concentrations, and a shift of the redox state of ascorbate to its oxidized form. In addition, ethylene-treated plants showed higher visual leaf area damage and greater reductions in the maximum efficiency of the PSII photochemistry than controls in response to heat stress or to a combination of heat and drought stress. These results demonstrate for the first time that airborne ethylene at concentrations similar to those found in polluted areas may reduce plant stress tolerance by altering, among other possible mechanisms, antioxidant defenses. © 2004 American Society of Plant Biologists.

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