Barba J., Curiel Yuste J., Poyatos R., Janssens I.A., Lloret F. (2016) Strong resilience of soil respiration components to drought-induced die-off resulting in forest secondary succession. Oecologia. 182: 27-41.EnlaceDoi: 10.1007/s00442-016-3567-8
How forests cope with drought-induced perturbations and how the dependence of soil respiration on environmental and biological drivers is affected in a warming and drying context are becoming key questions. The aims of this study were to determine whether drought-induced die-off and forest succession were reflected in soil respiration and its components and to determine the influence of climate on the soil respiration components. We used the mesh exclusion method to study seasonal variations in soil respiration (RS) and its components: heterotrophic (RH) and autotrophic (RA) [further split into fine root (RR) and mycorrhizal respiration (RM)] in a mixed Mediterranean forest where Scots pine (Pinus sylvestris L.) is undergoing a drought-induced die-off and is being replaced by holm oak (Quercus ilex L.). Drought-induced pine die-off was not reflected in RS nor in its components, which denotes a high functional resilience of the plant and soil system to pine die-off. However, the succession from Scots pine to holm oak resulted in a reduction of RH and thus in an important decrease of total respiration (RS was 36 % lower in holm oaks than in non-defoliated pines). Furthermore, RS and all its components were strongly regulated by soil water content-and-temperature interaction. Since Scots pine die-off and Quercus species colonization seems to be widely occurring at the driest limit of the Scots pine distribution, the functional resilience of the soil system over die-off and the decrease of RS from Scots pine to holm oak could have direct consequences for the C balance of these ecosystems. © 2016, Springer-Verlag Berlin Heidelberg.
Aguade D., Poyatos R., Gomez M., Oliva J., Martinez-Vilalta J. (2015) The role of defoliation and root rot pathogen infection in driving the mode of drought-related physiological decline in Scots pine (Pinus sylvestris L.). Tree Physiology. 35: 229-242.EnlaceDoi: 10.1093/treephys/tpv005
Drought-related tree die-off episodes have been observed in all vegetated continents. Despite much research effort, however, the multiple interactions between carbon starvation, hydraulic failure and biotic agents in driving tree mortality under field conditions are still not well understood. We analysed the seasonal variability of non-structural carbohydrates (NSCs) in four organs (leaves, branches, trunk and roots), the vulnerability to embolism in roots and branches, native embolism (percentage loss of hydraulic conductivity (PLC)) in branches and the presence of root rot pathogens in defoliated and non-defoliated individuals in a declining Scots pine (Pinus sylvestris L.) population in the NE Iberian Peninsula in 2012, which included a particularly dry and warm summer. No differences were observed between defoliated and non-defoliated pines in hydraulic parameters, except for a higher vulnerability to embolism at pressures below-2 MPa in roots of defoliated pines. No differences were found between defoliation classes in branch PLC. Total NSC (TNSC, soluble sugars plus starch) values decreased during drought, particularly in leaves. Defoliation reduced TNSC levels across tree organs, especially just before (June) and during (August) drought. Root rot infection by the fungal pathogen Onnia P. Karst spp. was detected but it did not appear to be associated to tree defoliation. However, Onnia infection was associated with reduced leaf-specific hydraulic conductivity and sapwood depth, and thus contributed to hydraulic impairment, especially in defoliated pines. Infection was also associated with virtually depleted root starch reserves during and after drought in defoliated pines. Moreover, defoliated and infected trees tended to show lower basal area increment. Overall, our results show the intertwined nature of physiological mechanisms leading to drought-induced mortality and the inherent difficulty of isolating their contribution under field conditions. © The Author 2015. Published by Oxford University Press. All rights reserved.
Aguade D., Poyatos R., Rosas T., Martinez-Vilalta J. (2015) Comparative drought responses of Quercus ilex L. and Pinus sylvestris L. In a montane forest undergoing a vegetation shift. Forests. 6: 2505-2529.EnlaceDoi: 10.3390/f6082505
Different functional and structural strategies to cope with water shortage exist both within and across plant communities. The current trend towards increasing drought in many regions could drive some species to their physiological limits of drought tolerance, potentially leading to mortality episodes and vegetation shifts. In this paper, we study the drought responses of Quercus ilex and Pinus sylvestris in a montane Mediterranean forest where the former species is replacing the latter in association with recent episodes of drought-induced mortality. Our aim was to compare the physiological responses to variations in soil water content (SWC) and vapor pressure deficit (VPD) of the two species when living together in a mixed stand or separately in pure stands, where the canopies of both species are completely exposed to high radiation and VPD. P. sylvestris showed typical isohydric behavior, with greater losses of stomatal conductance with declining SWC and greater reductions of stored non-structural carbohydrates during drought, consistent with carbon starvation being an important factor in the mortality of this species. On the other hand, Q. ilex trees showed a more anisohydric behavior, experiencing more negative water potentials and higher levels of xylem embolism under extreme drought, presumably putting them at higher risk of hydraulic failure. In addition, our results show relatively small changes in the physiological responses of Q. ilex in mixed vs. pure stands, suggesting that the current replacement of P. sylvestris by Q. ilex will continue. © 2015 by the authors.
Caceres M.D., Martinez-Vilalta J., Coll L., Llorens P., Casals P., Poyatos R., Pausas J.G., Brotons L. (2015) Coupling a water balance model with forest inventory data to predict drought stress: The role of forest structural changes vs. climate changes. Agricultural and Forest Meteorology. 213: 77-90.EnlaceDoi: 10.1016/j.agrformet.2015.06.012
Mechanistic water balance models can be used to predict soil moisture dynamics and drought stress in individual forest stands. Predicting current and future levels of plant drought stress is important not only at the local scale, but also at larger, landscape to regional, scales, because these are the management scales at which adaptation and mitigation strategies are implemented. To obtain reliable predictions of soil moisture and plant drought stress over large extents, water balance models need to be complemented with detailed information about the spatial variation of vegetation and soil attributes. We designed, calibrated and validated a water balance model that produces annual estimates of drought intensity and duration for all plant cohorts in a forest stand. Taking Catalonia (NE Spain) as a case study, we coupled this model with plot records from two Spanish forest inventories in which species identity, diameter and height of plant cohorts were available. Leaf area index of each plant cohort was estimated from basal area using species-specific relationships. Vertical root distribution for each species in each forest plot was estimated by determining the distribution that maximized transpiration in the model, given average climatic conditions, soil attributes and stand density. We determined recent trends (period 1980-2010) in drought stress for the main tree species in Catalonia; where forest growth and densification occurs in many areas as a result of rural abandonment and decrease of forest management. Regional increases in drought stress were detected for most tree species, although we found high variation in stress changes among individual forest plots. Moreover, predicted trends in tree drought stress were mainly due to changes in leaf area occurred between the two forest inventories rather than to climatic trends. We conclude that forest structure needs to be explicitly considered in assessments of plant drought stress patterns and trends over large geographic areas, and that forest inventories are useful sources of data provided that reasonably good estimates of soil attributes and root distribution are available. Our approach coupled with recent improvements in forest survey technologies may allow obtaining spatially continuous and precise assessments of drought stress. Further efforts are needed to calibrate drought-related demographic processes before water balance and drought stress estimates can be fully used for the accurate prediction of drought impacts. © 2015 Elsevier B.V.
Salmon Y., Torres-Ruiz J.M., Poyatos R., Martinez-Vilalta J., Meir P., Cochard H., Mencuccini M. (2015) Balancing the risks of hydraulic failure and carbon starvation: A twig scale analysis in declining Scots pine. Plant, Cell and Environment. : 0-0.EnlaceDoi: 10.1111/pce.12572
Understanding physiological processes involved in drought-induced mortality is important for predicting the future of forests and for modelling the carbon and water cycles. Recent research has highlighted the variable risks of carbon starvation and hydraulic failure in drought-exposed trees. However, little is known about the specific responses of leaves and supporting twigs, despite their critical role in balancing carbon acquisition and water loss. Comparing healthy (non-defoliated) and unhealthy (defoliated) Scots pine at the same site, we measured the physiological variables involved in regulating carbon and water resources. Defoliated trees showed different responses to summer drought compared with non-defoliated trees. Defoliated trees maintained gas exchange while non-defoliated trees reduced photosynthesis and transpiration during the drought period. At the branch scale, very few differences were observed in non-structural carbohydrate concentrations between health classes. However, defoliated trees tended to have lower water potentials and smaller hydraulic safety margins. While non-defoliated trees showed a typical response to drought for an isohydric species, the physiology appears to be driven in defoliated trees by the need to maintain carbon resources in twigs. These responses put defoliated trees at higher risk of branch hydraulic failure and help explain the interaction between carbon starvation and hydraulic failure in dying trees. Understanding the physiological responses of leaves to drought is crucial since they are the site of both photosynthesis and transpiration, and hence play key roles in balancing the risks of carbon starvation and hydraulic failure. Co-occurring healthy and unhealthy Scots pines showed different responses to summer drought: while healthy trees showed a typical response to drought for an isohydric species, atypical physiology in unhealthy trees appears to be driven by the need to maintain carbohydrate availability in needles and twigs. These responses put unhealthy trees at higher risk of branch hydraulic failure and help to explain the interaction between carbon-starvation and hydraulic failure in dying trees. © 2015 John Wiley & Sons Ltd.
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.EnlaceDoi: 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.
Martinez-Vilalta J., Poyatos R., Aguade D., Retana J., Mencuccini M. (2014) A new look at water transport regulation in plants. New Phytologist. : 0-0.EnlaceDoi: 10.1111/nph.12912
Plant function requires effective mechanisms to regulate water transport at a variety of scales. Here, we develop a new theoretical framework describing plant responses to drying soil, based on the relationship between midday and predawn leaf water potentials. The intercept of the relationship (Λ) characterizes the maximum transpiration rate per unit of hydraulic transport capacity, whereas the slope (σ) measures the relative sensitivity of the transpiration rate and plant hydraulic conductance to declining water availability. This framework was applied to a newly compiled global database of leaf water potentials to estimate the values of Λ and σ for 102 plant species. Our results show that our characterization of drought responses is largely consistent within species, and that the parameters Λ and σ show meaningful associations with climate across species. Parameter σ was ≤1 in most species, indicating a tight coordination between the gas and liquid phases of water transport, in which canopy transpiration tended to decline faster than hydraulic conductance during drought, thus reducing the pressure drop through the plant. The quantitative framework presented here offers a new way of characterizing water transport regulation in plants that can be used to assess their vulnerability to drought under current and future climatic conditions. © 2014 New Phytologist Trust.
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.EnlaceDoi: 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.
Poyatos R., Heinemeyer A., Ineson P., Evans J.G., Ward H.C., Huntley B., Baxter R. (2014) Environmental and Vegetation Drivers of Seasonal CO2 Fluxes in a Sub-arctic Forest-Mire Ecotone. Ecosystems. 17: 377-393.EnlaceDoi: 10.1007/s10021-013-9728-2
Unravelling the role of structural and environmental drivers of gross primary productivity (GPP) and ecosystem respiration (R eco) in highly heterogeneous tundra is a major challenge for the upscaling of chamber-based CO2 fluxes in Arctic landscapes. In a mountain birch woodland-mire ecotone, we investigated the role of LAI (and NDVI), environmental factors (microclimate, soil moisture), and microsite type across tundra shrub plots (wet hummocks, dry hummocks, dry hollows) and lichen hummocks, in controlling net ecosystem CO2 exchange (NEE). During a growing season, we measured NEE fluxes continuously, with closed dynamic chambers, and performed multiple fits (one for each 3-day period) of a simple light and temperature response model to hourly NEE data. Tundra shrub plots were largely CO2 sinks, as opposed to lichen plots, although fluxes were highly variable within microsite type. For tundra shrub plots, microsite type did not influence photosynthetic parameters but it affected basal (that is, temperature-normalized) ecosystem respiration (R 0). PAR-normalized photosynthesis (P 600) increased with air temperature and declined with increasing vapor pressure deficit. R 0 declined with soil moisture and showed an apparent increase with temperature, which may underlie a tight link between GPP and R eco. NDVI was a good proxy for LAI, maximum P 600 and maximum R 0 of shrub plots. Cumulative CO2 fluxes were strongly correlated with LAI (NDVI) but we observed a comparatively low GPP/LAI in dry hummocks. Our results broadly agree with the reported functional convergence across tundra vegetation, but here we show that the role of decreased productivity in transition zones and the influence of temperature and water balance on seasonal CO2 fluxes in sub-Arctic forest-mire ecotones cannot be overlooked. © 2013 Springer Science+Business Media New York.
Sus O., Poyatos R., Barba J., Carvalhais N., Llorens P., Williams M., Vilalta J.M. (2014) Time variable hydraulic parameters improve the performance of a mechanistic stand transpiration model. A case study of Mediterranean Scots pine sap flow data assimilation. Agricultural and Forest Meteorology. 198: 168-180.EnlaceDoi: 10.1016/j.agrformet.2014.08.009
Tree transpiration is regulated by short-term physiological adjustments and long-term shifts in hydraulic architecture in response to fluctuating evaporative demand and water supply. Despite the tight interdependence of plant water loss and carbon uptake and its crucial implications for plant growth and survival under drought conditions, the underlying mechanisms remain incompletely represented in most state-of-the-art mechanistic models. Important process information is resolved in tree transpiration (sap flow) data, which are the measurable outcome of water transport through the soil-plant-atmosphere continuum under variable environmental conditions. Here, we assimilated sap flow data measured in two Scots pine stands from climatically contrasting sites - one of which experiencing a strong drought during the study period - in NE Spain into a process-based ecophysiological model (SPA) using the Ensemble Kalman Filter (EnKF) in order to: (1) distinguish differences in hydraulic characteristics between sites and between healthy and defoliated individuals within a site; (2) identify possible structural model deficiencies, particularly regarding temporal changes in plant hydraulic conductance which the model assumes constant; and (3) derive implications for gross photosynthesis and carbon cycling. In terms of stomatal control, the assimilation of sap flow data into SPA showed a more conservative water use under dry conditions. Time-varying plant conductivity substantially improved model performance under severe drought, while seasonally varying capacitance and stomatal efficiency only resulted in marginal improvements. Not accounting for this seasonal variability would translate into a 30-60% overestimation of modelled GPP during drought. Our results suggest that an explicit representation of mechanisms leading to temporal changes in hydraulic conductivity (i.e., xylem embolism) is required for models to reproduce tree functioning under extreme drought.
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