Sperlich D., Chang C.T., Penuelas J., Gracia C., Sabate S. (2015) Seasonal variability of foliar photosynthetic and morphological traits and drought impacts in a Mediterranean mixed forest. Tree Physiology. 35: 501-520.EnllaçDoi: 10.1093/treephys/tpv017
The Mediterranean region is a hot spot of climate change vulnerable to increased droughts and heat waves. Scaling carbon fluxes from leaf to landscape levels is particularly challenging under drought conditions. We aimed to improve the mechanistic understanding of the seasonal acclimation of photosynthesis and morphology in sunlit and shaded leaves of four Mediterranean trees (Quercus ilex L., Pinus halepensis Mill., Arbutus unedo L. and Quercus pubescens Willd.) under natural conditions. Vc,max and Jmax were not constant, and mesophyll conductance was not infinite, as assumed in most terrestrial biosphere models, but varied significantly between seasons, tree species and leaf position. Favourable conditions in winter led to photosynthetic recovery and growth in the evergreens. Under moderate drought, adjustments in the photo/biochemistry and stomatal/mesophyllic diffusion behaviour effectively protected the photosynthetic machineries. Severe drought, however, induced early leaf senescence mostly in A. unedo and Q. pubescens, and significantly increased leaf mass per area in Q. ilex and P. halepensis. Shaded leaves had lower photosynthetic potentials but cushioned negative effects during stress periods. Species-specificity, seasonal variations and leaf position are key factors to explain vegetation responses to abiotic stress and hold great potential to reduce uncertainties in terrestrial biosphere models especially under drought conditions. © The Author 2015. Published by Oxford University Press.
Sánchez-Costa E., Poyatos R., Sabaté S. (2015) Contrasting growth and water use strategies in four co-occurring Mediterranean tree species revealed by concurrent measurements of sap flow and stem diameter variations. Agricultural and Forest Meteorology. 207: 24-37.EnllaçDoi: 10.1016/j.agrformet.2015.03.012
Drought limits tree water use and growth of Mediterranean trees. However, growth and water use strategies are rarely addressed simultaneously across species and drought conditions. Here, we investigate the link between stem diameter variations and sap flow in four co-existing Mediterranean trees (Pinus halepensis Mill., Quercus pubescens Willd., Quercus ilex L. and Arbutus unedo L.), under relatively wet (2011) and dry (2012) conditions. Continuous stem diameter variations were converted to basal area increment (BAI) and de-trended to estimate tree water deficit (δW), an indicator of stem hydration. P. halepensis and Q. pubescens showed the most and the least conservative sap flow density (JS) regulation under drought, respectively, with Q. ilex and A.unedo showing intermediate drought responses. All species, except A. unedo, showed some between-year variability in the environmental control of JS. Seasonal stem shrinkage in response to drought (i.e., increasing δW) and subsequent trunk rehydration after rainfall (i.e., decreasing δW) occurred in all species. Vapor pressure deficit (VPD) and soil moisture (θ) interacted to determine seasonal variation in δW. Interestingly, in the dry year, 2012, more species-specific differences were found in the responses of δW to θ and VPD. Across species, JS and δW began to decline at similar soil moisture thresholds, underpinning the tight link between JS and δW under varying drought conditions. Annual BAI decreased proportionally more than tree-level transpiration (JT) between the wet (2011) and the dry (2012) year, hence growth-based WUE (WUEBAI=BAI/JT) decreased for all species, albeit less acutely for P. halepensis. Overall, despite their contrasting leaf habit and wood type, the studied Mediterranean tree species show coordinated responses of transpiration, water storage dynamics and growth-based WUE which allow them to cope with seasonal and interannual drought. © 2015 Elsevier B.V.
Audsley E., Trnka M., Sabate S., Maspons J., Sanchez A., Sandars D., Balek J., Pearn K. (2014) Interactively modelling land profitability to estimate European agricultural and forest land use under future scenarios of climate, socio-economics and adaptation. Climatic Change. : 0-0.EnllaçDoi: 10.1007/s10584-014-1164-6
Studies of climate change impacts on agricultural land use generally consider sets of climates combined with fixed socio-economic scenarios, making it impossible to compare the impact of specific factors within these scenario sets. Analysis of the impact of specific scenario factors is extremely difficult due to prohibitively long run-times of the complex models. This study produces and combines metamodels of crop and forest yields and farm profit, derived from previously developed very complex models, to enable prediction of European land use under any set of climate and socio-economic data. Land use is predicted based on the profitability of the alternatives on every soil within every 10' grid across the EU. A clustering procedure reduces 23,871 grids with 20+ soils per grid to 6,714 clusters of common soil and climate. Combined these reduce runtime 100 thousand-fold. Profit thresholds define land as intensive agriculture (arable or grassland), extensive agriculture or managed forest, or finally unmanaged forest or abandoned land. The demand for food as a function of population, imports, food preferences and bioenergy, is a production constraint, as is irrigation water available. An iteration adjusts prices to meet these constraints. A range of measures are derived at 10' grid-level such as diversity as well as overall EU production. There are many ways to utilise this ability to do rapid What-If analysis of both impact and adaptations. The paper illustrates using two of the 5 different GCMs (CSMK3, HADGEM with contrasting precipitation and temperature) and two of the 4 different socio-economic scenarios ("We are the world", "Should I stay or should I go" which have contrasting demands for land), exploring these using two of the 13 scenario parameters (crop breeding for yield and population) . In the first scenario, population can be increased by a large amount showing that food security is far from vulnerable. In the second scenario increasing crop yield shows that it improves the food security problem. © 2014 Springer Science+Business Media Dordrecht.
Chang C.T., Sabaté S., Sperlich D., Poblador S., Sabater F., Gracia C. (2014) Does soil moisture overrule temperature dependence of soil respiration in Mediterranean riparian forests?. Biogeosciences. 11: 6173-6185.EnllaçDoi: 10.5194/bg-11-6173-2014
Soil respiration (SR) is a major component of ecosystems' carbon cycles and represents the second largest CO2 flux in the terrestrial biosphere. Soil temperature is considered to be the primary abiotic control on SR, whereas soil moisture is the secondary control factor. However, soil moisture can become the dominant control on SR in very wet or dry conditions. Determining the trigger that makes soil moisture as the primary control factor of SR will provide a deeper understanding on how SR changes under the projected future increase in droughts. Specific objectives of this study were (1) to investigate the seasonal variations and the relationship between SR and both soil temperature and moisture in a Mediterranean riparian forest along a groundwater level gradient; (2) to determine soil moisture thresholds at which SR is controlled by soil moisture rather than by temperature; (3) to compare SR responses under different tree species present in a Mediterranean riparian forest (Alnus glutinosa, Populus nigra and Fraxinus excelsior). Results showed that the heterotrophic soil respiration rate, groundwater level and 30 cm integral soil moisture (SM30) decreased significantly from the riverside moving uphill and showed a pronounced seasonality. SR rates showed significant differences between tree species, with higher SR for P. nigra and lower SR for A. glutinosa. The lower threshold of soil moisture was 20 and 17% for heterotrophic and total SR, respectively. Daily mean SR rate was positively correlated with soil temperature when soil moisture exceeded the threshold, with Q10 values ranging from 1.19 to 2.14; nevertheless, SR became decoupled from soil temperature when soil moisture dropped below these thresholds. © 2014 Author(s).
Moran-Lopez T., Poyatos R., Llorens P., Sabate S. (2014) Effects of past growth trends and current water use strategies on scots pine and pubescent oak drought sensitivity. European Journal of Forest Research. 133: 369-382.EnllaçDoi: 10.1007/s10342-013-0768-0
Drought-induced decline is affecting Pinus sylvestris populations in southern Europe, with very little impact on the more drought-tolerant Quercus pubescens. Although multiple studies have investigated interspecific differences in water use and growth strategies, the link between these two processes and how they vary within drought-exposed populations remains poorly understood. Here, we analysed tree ring and sap flow data from P. sylvestris and Q. pubescens stands in the Pyrenees in order to (1) evaluate differences in climate-growth responses among species, (2) disentangle the role of past growth trends and water use strategies in individual trees drought sensitivity and (3) assess whether such intraspecific patterns vary between species. Both species have suffered recent climatic constraints related to increased aridity. However, the effects of past growth trends and current water use traits on drought sensitivity varied among them. Initially, fast-growing 'drought-sensitive' pines displayed a higher gas exchange potential but were more sensitive to evaporative demand and soil moisture. They also showed lower water use efficiency for growth (WUEBAI) and current growth decline. In contrast, initially, slow-growing 'drought-tolerant' pines showed the opposite water use traits and currently maintain the highest growth rates. In comparison, neither current WUEBAI nor recent growth trends varied across Q. pubescens climate-growth groups. Nonetheless, 'drought-sensitive' oaks showed the lowest gas exchange potential and the highest growth rates under milder conditions. Our results show a strong effect of past growth trends and current water use strategies on tree resilience to increased aridity, which is more evident in P. sylvestris. © Springer-Verlag Berlin Heidelberg 2013.
Santos X., Mateos E., Bros V., Brotons L., De Mas E., Herraiz J.A., Herrando S., Mino A., Olmo-Vidal J.M., Quesada J., Ribes J., Sabate S., Sauras-Yera T., Serra A., Ramon Vallejo V., Vinolas A. (2014) Is response to fire influenced by dietary specialization and mobility? A comparative study with multiple animal assemblages. PLoS ONE. 9: 0-0.EnllaçDoi: 10.1371/journal.pone.0088224
Fire is a major agent involved in landscape transformation and an indirect cause of changes in species composition. Responses to fire may vary greatly depending on life histories and functional traits of species. We have examined the taxonomic and functional responses to fire of eight taxonomic animal groups displaying a gradient of dietary and mobility patterns: Gastropoda, Heteroptera, Formicidae, Coleoptera, Araneae, Orthoptera, Reptilia and Aves. The fieldwork was conducted in a Mediterranean protected area on 3 sites (one unburnt and two burnt with different postfire management practices) with five replicates per site. We collected information from 4606 specimens from 274 animal species. Similarity in species composition and abundance between areas was measured by the Bray-Curtis index and ANOSIM, and comparisons between animal and plant responses by Mantel tests. We analyze whether groups with the highest percentage of omnivorous species, these species being more generalist in their dietary habits, show weak responses to fire (i.e. more similarity between burnt and unburnt areas), and independent responses to changes in vegetation. We also explore how mobility, i.e. dispersal ability, influences responses to fire. Our results demonstrate that differences in species composition and abundance between burnt and unburnt areas differed among groups. We found a tendency towards presenting lower differences between areas for groups with higher percentages of omnivorous species. Moreover, taxa with a higher percentage of omnivorous species had significantly more independent responses of changes in vegetation. High- (e.g. Aves) and low-mobility (e.g. Gastropoda) groups had the strongest responses to fire (higher R scores of the ANOSIM); however, we failed to find a significant general pattern with all the groups according to their mobility. Our results partially support the idea that functional traits underlie the response of organisms to environmental changes caused by fire. © 2014 Santos et al.
Sperlich D., Chang C.T., Penuelas J., Gracia C., Sabate S. (2014) Foliar photochemical processes and carbon metabolism under favourable and adverse winter conditions in a Mediterranean mixed forest, Catalonia (Spain). Biogeosciences. 11: 5657-5674.EnllaçDoi: 10.5194/bg-11-5657-2014
Evergreen trees in the Mediterranean region must cope with a wide range of environmental stresses from summer drought to winter cold. The mildness of Mediterranean winters can periodically lead to favourable environmental conditions above the threshold for a positive carbon balance, benefitting evergreen woody species more than deciduous ones. The comparatively lower solar energy input in winter decreases the foliar light saturation point. This leads to a higher susceptibility to photoinhibitory stress especially when chilly (< 12 °C) or freezing temperatures (< 0 °C) coincide with clear skies and relatively high solar irradiances. Nonetheless, the advantage of evergreen species that are able to photosynthesize all year round where a significant fraction can be attributed to winter months, compensates for the lower carbon uptake during spring and summer in comparison to deciduous species. We investigated the ecophysiological behaviour of three co-occurring mature evergreen tree species (Quercus ilex L., Pinus halepensis Mill., and Arbutus unedo L.). Therefore, we collected twigs from the field during a period of mild winter conditions and after a sudden cold period. After both periods, the state of the photosynthetic machinery was tested in the laboratory by estimating the foliar photosynthetic potential with CO2 response curves in parallel with chlorophyll fluorescence measurements. The studied evergreen tree species benefited strongly from mild winter conditions by exhibiting extraordinarily high photosynthetic potentials. A sudden period of frost, however, negatively affected the photosynthetic apparatus, leading to significant decreases in key physiological parameters such as the maximum carboxylation velocity (Vc, max), the maximum photosynthetic electron transport rate (Jmax), and the optimal fluorometric quantum yield of photosystem II (Fv/Fm). The responses of Vc, max and Jmax were highly species specific, with Q. ilex exhibiting the highest and P. halepensis the lowest reductions. In contrast, the optimal fluorometric quantum yield of photosystem II (Fv/Fm) was significantly lower in <i>A. unedo</i> after the cold period. The leaf position played an important role in Q. ilex showing a stronger winter effect on sunlit leaves in comparison to shaded leaves. Our results generally agreed with the previous classifications of photoinhibition-tolerant (P. halepensis) and photoinhibition-avoiding (Q. ilex) species on the basis of their susceptibility to dynamic photoinhibition, whereas A. unedo was the least tolerant to photoinhibition, which was chronic in this species. Q. ilex and P. halepensis seem to follow contrasting photoprotective strategies. However, they seemed equally successful under the prevailing conditions exhibiting an adaptive advantage over A. unedo. These results show that our understanding of the dynamics of interspecific competition in Mediterranean ecosystems requires consideration of the physiological behaviour during winter which may have important implications for long-term carbon budgets and growth trends.
Zhou S., Medlyn B., Sabaté S., Sperlich D., Colin Prentice I. (2014) Short-term water stress impacts on stomatal, mesophyll and biochemical limitations to photosynthesis differ consistently among tree species from contrasting climates. Tree Physiology. 34: 1035-1046.EnllaçDoi: 10.1093/treephys/tpu072
Predicting the large-scale consequences of drought in contrasting environments requires that we understand how drought effects differ among species originating from those environments. A previous meta-analysis of published experiments suggested that the effects of drought on both stomatal and non-stomatal limitations to photosynthesis may vary consistently among species from different hydroclimates. Here, we explicitly tested this hypothesis with two short-term water stress experiments on congeneric mesic and xeric species. One experiment was run in Australia using Eucalyptus species and the second was run in Spain using Quercus species as well as two more mesic species. In each experiment, plants were grown under moist conditions in a glasshouse, then deprived of water, and gas exchange was monitored. The stomatal response was analysed with a recently developed stomatal model, whose single parameter g1 represents the slope of the relationship between stomatal conductance and photosynthesis. The non-stomatal response was partitioned into effects on mesophyll conductance (gm), the maximum Rubisco activity (Vcmax) and the maximum electron transport rate (Jmax). We found consistency among the drought responses of g1, gm, Vcmax and Jmax, suggesting that drought imposes limitations on Rubisco activity and RuBP regeneration capacity concurrently with declines in stomatal and mesophyll conductance. Within each experiment, the more xeric species showed relatively high g1 under moist conditions, low drought sensitivity of g1, gm, Vcmax and Jmax, and more negative values of the critical pre-dawn water potential at which Vcmax declines most steeply, compared with the more mesic species. These results indicate adaptive interspecific differences in drought responses that allow xeric tree species to continue transpiration and photosynthesis for longer during periods without rain. © The Author 2014. Published by Oxford University Press. All rights reserved.
Serra-Diaz J.M., Keenan T.F., Ninyerola M., Sabate S., Gracia C., Lloret F. (2013) Geographical patterns of congruence and incongruence between correlative species distribution models and a process-based ecophysiological growth model. Journal of Biogeography. 40: 1928-1938.EnllaçDoi: 10.1111/jbi.12142
Aim: Our aim was to map the climate dependence of tree species distributions (probability of occurrence) and forest growth (net primary productivity) by comparing the congruence and incongruence between correlative and process-based modelling approaches. Location: Iberian Peninsula, south-western Europe. Methods: We used forest inventory data for three widespread tree species (Quercus ilex, Pinus halepensis and Pinus sylvestris) to model climatic suitability with an ensemble of seven correlative species distribution models (using biomod). We then simulated forest net primary productivity (NPP) as a surrogate of forest growth for forests of each species using an ecophysiological process-based model (gotilwa+) along a gradient of climatic suitability. The spatial distribution of the growth estimates was then compared with that of the suitability estimates, and robust regression was used to classify regions in terms of model congruence. Results: Quercus ilex and P. sylvestris both showed a positive relationship between forest NPP and climatic suitability. The main discrepancies were found in the north of the peninsula, where there was high potential forest growth but low climate suitability. Low forest-growth estimates in areas of high suitability only appeared for P. sylvestris in southern montane regions. Pinus halepensis always showed a negative relationship between estimated growth and climatic suitability. The analysis of other ecophysiological parameters (mean leaf life and leaf area index) suggests that this tree species has different physiological strategies that allow differential growth rates in areas of low suitability. Main conclusions: We found that the relationship between estimated growth and distribution varies strongly in different areas and species. Mapping the incongruences between the predicted climatic suitability and growth allowed us to identify regions where other factors (e.g. biotic interactions) may be more significant than the physiological limits on growth. We show that new insights into species distributions can be gained from mapping the differences between correlative and process-based models. © 2013 John Wiley & Sons Ltd.
Sperlich D, Sanchez E, Chang C, Gracia C, Peñuelas J, Sabaté S (2012) Seasonal photosynthetic behaviour and drought. Responses of Mediterranean tree species. GreencyclesII Mini-conference MC3, "Evaluation of Earth system models using modern and palaeoobservations". Clare College, Cambridge. 24-25 September.
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