Barbeta A., Camarero J.J., Sangüesa-Barreda G., Muffler L., Peñuelas J. (2019) Contrasting effects of fog frequency on the radial growth of two tree species in a Mediterranean-temperate ecotone. Agricultural and Forest Meteorology. 264: 297-308.EnllaçDoi: 10.1016/j.agrformet.2018.10.020
The performance and persistence of rear-edge tree populations are relevant issues for conserving biodiversity because these stands harbor high intraspecific biodiversity and play a key role during periods of climate change. The occurrence of these populations is associated with the influence of heterogeneous topography, creating suitable refugia with regionally rare environmental conditions. Climate is changing at a global-scale, but little is known about the long-term impact on local climatic singularities and the associated taxa. We analyzed tree-ring growth chronologies of the two species (Fagus sylvatica and Quercus ilex) forming the evergreen-deciduous forest ecotone, constitutive of the rear-edge of F. sylvatica distribution. The study area is a coastal range with frequent fog immersion, which has been hypothesized to favor the persistence of F. sylvatica in Mediterranean peninsulas. We analyzed the long-term effect of fog on tree growth along a topographical gradient and the sensitivity of growth to rainfall and temperature. The annual number of foggy days has decreased by 62% over the last four decades, concomitant with increasing temperatures. Fog frequency was a relevant factor determining tree growth; fog during summer had positive effects on F. sylvatica growth mainly through a temperature buffering effect. The positive effect of fog on the growth of Q. ilex, however, was likely caused by a collinearity with rainfall. Q. ilex growth was less sensitive to climate than F. sylvatica, but growth of both species was enhanced by a positive early-summer water balance. Our results indicate that a decrease in fog frequency and an increase in temperature may generally benefit Q. ilex in this forest ecotone. Although future changes in rainfall and temperature matter most for the fate of rear-edge tree populations, local climatic singularities such as fog should also be considered. Those can have complementary effects that can swing the balance in ecotones and rear-edge tree populations such as those studied here. © 2018 Elsevier B.V.
Liu D., Ogaya R., Barbeta A., Yang X., Peñuelas J. (2018) Long-term experimental drought combined with natural extremes accelerate vegetation shift in a Mediterranean holm oak forest. Environmental and Experimental Botany. 151: 1-11.EnllaçDoi: 10.1016/j.envexpbot.2018.02.008
Increasing drought combined with natural extremes are expected to accelerate forest die-off and shifts in vegetation in the Mediterranean Basin. However, fewer studies have explored these climate-driven changes in forest ecosystems. A long-term (17-year) experimental drought (−30% precipitation) was established in a Mediterranean holm oak forest with high (H) and low (L) canopies to determine the changes in stem mortality, recruitment and composition shifts. Experimental drought increased annual stem mortality rate at the community level for both H- and L-canopies. Natural drought amplified the effects of experimental drought on stem mortality at the community level and of Q. ilex for H- and L-canopies. The timescales of natural drought, however, varied substantially with canopy types and species, with shorter timescales in L- than H-canopy and for Q. ilex than P. latifolia. Furthermore, experimental drought combined with natural extremes amplified the increases in stem mortality and decreases in growth for L-canopy. Contrasting responses between Q. ilex and P. latifolia for the relative in abundance and growth were observed in L-canopy and drought treatment reinforced the vegetation shift favoring P. latifolia. These findings suggest continuous drought regimes accelerated a vegetation shift, implying potential consequences for the functions and services for water-limited forest ecosystems. © 2018 Elsevier B.V.
Barbeta A., Peñuelas J. (2017) Relative contribution of groundwater to plant transpiration estimated with stable isotopes. Scientific Reports. 7: 0-0.EnllaçDoi: 10.1038/s41598-017-09643-x
Water stored underground in the saturated and subsurface zones below the soil are important sources of water for plants in water-limited ecosystems. The presence of deep-rooted plants worldwide, however, suggests that the use of groundwater is not restricted to arid and seasonally dry ecosystems. We compiled the available data (71 species) on the relative contribution of groundwater to plant water estimated using stable isotopes and mixing models, which provided information about relative groundwater use, and analyzed their variation across different climates, seasons, plant types, edaphic conditions, and landscape positions. Plant use of groundwater was more likely at sites with a pronounced dry season, and represented on average 49 per cent of transpired water in dry seasons and 28 per cent in wet seasons. The relative contribution of groundwater to plant-water uptake was higher on rocky substrates (saprolite, fractured bedrock), which had reduced groundwater uptake when this source was deep belowground. In addition, we found that the connectivity between groundwater pools and plant water may be quantitatively larger and more widespread than reported by recent global estimations based on isotopic averaged values. Earth System Models should account for the feedbacks between transpiration and groundwater recharge. © 2017 The Author(s).
Rivas-Ubach A., Barbeta A., Sardans J., Guenther A., Ogaya R., Oravec M., Urban O., Peñuelas J. (2016) Topsoil depth substantially influences the responses to drought of the foliar metabolomes of Mediterranean forests. Perspectives in Plant Ecology, Evolution and Systematics. 21: 41-54.EnllaçDoi: 10.1016/j.ppees.2016.06.001
The upper soil provides support, water, and nutrients to terrestrial plants and is therefore crucial for forest dynamics. We hypothesised that a tree's metabolic activity (and therefore its metabolome; the total set of metabolites) would be affected by both the depth of upper soil layers and water availability. We sampled leaves for stoichiometric and metabolomic analyses once per season from differently sized Quercus ilex trees under natural and experimental drought conditions representing the likely conditions in the coming decades). Although the metabolomes varied according to tree size, smaller trees did not show higher concentrations of biomarker metabolites related to drought stress. However, the effect of the drought treatment on the metabolomes was greatest for small trees growing in shallow soils. Our results suggest that tree size is more dependent on the depth of the upper soil, which indirectly affects a tree's metabolome, rather than on the moisture content in the upper soil. Metabolomic profiling of Q. ilex supports our finding that water availability in the upper soil is not necessarily correlated with tree size. The higher impact of drought on trees growing in shallower soils nevertheless indicates that any increase in the frequency, intensity, and duration of drought - as has been projected for the Mediterranean Basin and other areas - would affect small trees most. Metabolomics has proved to be a useful means for investigating the links between plant metabolism and environmental conditions. © 2016.
Sperlich D., Barbeta A., Ogaya R., Sabaté S., Peñuelas J. (2016) Balance between carbon gain and loss under long-term drought: Impacts on foliar respiration and photosynthesis in Quercus ilex L. Journal of Experimental Botany. 67: 821-833.EnllaçDoi: 10.1093/jxb/erv492
Terrestrial carbon exchange is a key process of the global carbon cycle consisting of a delicate balance between photosynthetic carbon uptake and respiratory release. We have, however, a limited understanding how long-term decreases in precipitation induced by climate change affect the boundaries and mechanisms of photosynthesis and respiration. We examined the seasonality of photosynthetic and respiratory traits and evaluated the adaptive mechanism of the foliar carbon balance of Quercus ilex L. experiencing a long-term rainfall-exclusion experiment. Day respiration (R d) but not night respiration (R n) was generally higher in the drought treatment leading to an increased R d/R n ratio. The limitation of mesophyll conductance (g m) on photosynthesis was generally stronger than stomatal limitation (g s) in the drought treatment, reflected in a lower g m/g s ratio. The peak photosynthetic activity in the drought treatment occurred in an atypical favourable summer in parallel with lower R d/R n and higher g m/g s ratios. The plant carbon balance was thus strongly improved through: (i) higher photosynthetic rates induced by g m; and (ii) decreased carbon losses mediated by R d. Interestingly, photosynthetic potentials (V c,max, J max, and TPU) were not affected by the drought treatment, suggesting a dampening effect on the biochemical level in the long term. In summary, the trees experiencing a 14-year-long drought treatment adapted through higher plasticity in photosynthetic and respiratory traits, so that eventually the atypical favourable growth period was exploited more efficiently. © 2015 The Author 2015. Published by Oxford University Press on behalf of the Society for Experimental Biology.
Liu D., Ogaya R., Barbeta A., Yang X., Peñuelas J. (2015) Contrasting impacts of continuous moderate drought and episodic severe droughts on the aboveground-biomass increment and litterfall of three coexisting Mediterranean woody species. Global Change Biology. 21: 4196-4209.EnllaçDoi: 10.1111/gcb.13029
Climate change is predicted to increase the aridity in the Mediterranean Basin and severely affect forest productivity and composition. The responses of forests to different timescales of drought, however, are still poorly understood because extreme and persistent moderate droughts can produce nonlinear responses in plants. We conducted a rainfall-manipulation experiment in a Mediterranean forest dominated by Quercus ilex, Phillyrea latifolia, and Arbutus unedo in the Prades Mountains in southern Catalonia from 1999 to 2014. The experimental drought significantly decreased forest aboveground-biomass increment (ABI), tended to increase the litterfall, and decreased aboveground net primary production throughout the 15 years of the study. The responses to the experimental drought were highly species-specific. A. unedo suffered a significant reduction in ABI, Q. ilex experienced a decrease during the early experiment (1999-2003) and in the extreme droughts of 2005-2006 and 2011-2012, and P. latifolia was unaffected by the treatment. The drought treatment significantly increased branch litterfall, especially in the extremely dry year of 2011, and also increased overall leaf litterfall. The drought treatment reduced the fruit production of Q. ilex, which affected seedling recruitment. The ABIs of all species were highly correlated with SPEI in early spring, whereas the branch litterfalls were better correlated with summer SPEIs and the leaf and fruit litterfalls were better correlated with autumn SPEIs. These species-specific responses indicated that the dominant species (Q. ilex) could be partially replaced by the drought-resistant species (P. latifolia). However, the results of this long-term study also suggest that the effect of drought treatment has been dampened over time, probably due to a combination of demographic compensation, morphological and physiological acclimation, and epigenetic changes. However, the structure of community (e.g., species composition, dominance, and stand density) may be reordered when a certain drought threshold is reached. © 2015 John Wiley & Sons Ltd.
Martin-Gomez P., Barbeta A., Voltas J., Penuelas J., Dennis K., Palacio S., Dawson T.E., Ferrio J.P. (2015) Isotope-ratio infrared spectroscopy: A reliable tool for the investigation of plant-water sources?. New Phytologist. : 0-0.EnllaçDoi: 10.1111/nph.13376
Summary: Stable isotopes are extensively used as tracers for the study of plant-water sources. Isotope-ratio infrared spectroscopy (IRIS) offers a cheaper alternative to isotope-ratio mass spectroscopy (IRMS), but its use in studying plant and soil water is limited by the spectral interference caused by organic contaminants. Here, we examine two approaches to cope with contaminated samples in IRIS: on-line oxidation of organic compounds (MCM) and post-processing correction. We assessed these methods compared to IRMS across 136 samples of xylem and soil water, and a set of ethanol- and methanol-water mixtures. A post-processing correction significantly improved IRIS accuracy in both natural samples and alcohol dilutions, being effective with concentrations up to 8% of ethanol and 0.4% of methanol. MCM outperformed the post-processing correction in removing methanol interference, but did not effectively remove interference for high concentrations of ethanol. By using both approaches, IRIS can overcome with reasonable accuracy the analytical uncertainties associated with most organic contaminants found in soil and xylem water. We recommend the post-processing correction as the first choice for analysis of samples of unknown contamination. Nevertheless, MCM can be more effective for evaluating samples containing contaminants responsible for strong spectral interferences at low concentrations, such as methanol. © 2015 New Phytologist Trust.
Ogaya R., Barbeta A., Basnou C., Penuelas J. (2015) Satellite data as indicators of tree biomass growth and forest dieback in a Mediterranean holm oak forest. Annals of Forest Science. 72: 135-144.EnllaçDoi: 10.1007/s13595-014-0408-y
• Context: In the framework of climate change, decreased tree growth and enhanced mortality induced by hot and dry conditions are increasing in many forests around the world, and particularly in Mediterranean forests.• Aims: Our aim was to estimate tree growth and mortality in a Mediterranean holm oak forest, using remote sensing data from MODIS.• Methods: We monitored annual increases of aboveground biomass by measuring tree basal area, and we determined tree mortality by counting dead stems. We analyzed the relationships between forest growth and mortality with mean annual values of some MODIS products and meteorological data.• Results: Mortality and increases of aboveground biomass correlated well with precipitation, September standardized precipitation/evapotranspiration indices (SPEI), and some MODIS products such as NDVI and enhanced vegetation index EVI. Other MODIS products such as gross primary production (GPP) and net photosynthesis, however, showed no clear relationship with tree mortality or measured increases of biomass.• Conclusion: The MODIS products as proxies of ecosystemic productivity (gross primary productivity, net photosynthesis) were weakly correlated with biomass increase, and did not reflect the mortality following the drought of autumn 2011. Nevertheless, NDVI and EVI were efficient indicators of forest productivity and dieback. © 2014, INRA and Springer-Verlag France.
Barbeta A., Mejia-Chang M., Ogaya R., Voltas J., Dawson T.E., Penuelas J. (2014) The combined effects of a long-term experimental drought and an extreme drought on the use of plant-water sources in a Mediterranean forest. Global Change Biology. : 0-0.EnllaçDoi: 10.1111/gcb.12785
Vegetation in water-limited ecosystems relies strongly on access to deep water reserves to withstand dry periods. Most of these ecosystems have shallow soils over deep groundwater reserves. Understanding the functioning and functional plasticity of species-specific root systems and the patterns of or differences in the use of water sources under more frequent or intense droughts is therefore necessary to properly predict the responses of seasonally dry ecosystems to future climate. We used stable isotopes to investigate the seasonal patterns of water uptake by a sclerophyll forest on sloped terrain with shallow soils. We assessed the effect of a long-term experimental drought (12 years) and the added impact of an extreme natural drought that produced widespread tree mortality and crown defoliation. The dominant species, Quercus ilex, Arbutus unedo and Phillyrea latifolia, all have dimorphic root systems enabling them to access different water sources in space and time. The plants extracted water mainly from the soil in the cold and wet seasons but increased their use of groundwater during the summer drought. Interestingly, the plants subjected to the long-term experimental drought shifted water uptake toward deeper (10-35 cm) soil layers during the wet season and reduced groundwater uptake in summer, indicating plasticity in the functional distribution of fine roots that dampened the effect of our experimental drought over the long term. An extreme drought in 2011, however, further reduced the contribution of deep soil layers and groundwater to transpiration, which resulted in greater crown defoliation in the drought-affected plants. This study suggests that extreme droughts aggravate moderate but persistent drier conditions (simulated by our manipulation) and may lead to the depletion of water from groundwater reservoirs and weathered bedrock, threatening the preservation of these Mediterranean ecosystems in their current structures and compositions.
Ogaya R., Llusia J., Barbeta A., Asensio D., Liu D., Alessio G.A., Penuelas J. (2014) Foliar CO2 in a holm oak forest subjected to 15 years of climate change simulation. Plant Science. 226: 101-107.EnllaçDoi: 10.1016/j.plantsci.2014.06.010
A long-term experimental drought to simulate future expected climatic conditions for Mediterranean forests, a 15% decrease in soil moisture for the following decades, was conducted in a holm oak forest since 1999. Net photosynthetic rate, stomatal conductance and leaf water potential were measured from 1999 to 2013 in Quercus ilex and Phillyrea latifolia, two co-dominant species of this forest. These measurements were performed in four plots, two of them received the drought treatment and the two other plots were control plots. The three studied variables decreased with increases in VPD and decreases in soil moisture in both species, but the decrease of leaf water potential during summer drought was larger in P. latifolia, whereas Q. ilex reached higher net photosynthetic rates and stomatal conductance values during rainy periods than P. latifolia. The drought treatment decreased ca. 8% the net photosynthetic rates during the overall studied period in both Q. ilex and P. latifolia, whereas there were just non-significant trends toward a decrease in leaf water potential and stomatal conductance induced by drought treatment. Future drier climate may lead to a decrease in the carbon balance of Mediterranean species, and some shrub species well resistant to drought could gain competitive advantage relative to Q. ilex, currently the dominant species of this forest. © 2014 Elsevier Ireland Ltd.
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