Poyatos R., Sus O., Badiella L., Mencuccini M., Martínez-Vilalta J. (2018) Gap-filling a spatially explicit plant trait database: Comparing imputation methods and different levels of environmental information. Biogeosciences. 15: 2601-2617.EnlaceDoi: 10.5194/bg-15-2601-2018
The ubiquity of missing data in plant trait databases may hinder trait-based analyses of ecological patterns and processes. Spatially explicit datasets with information on intraspecific trait variability are rare but offer great promise in improving our understanding of functional biogeography. At the same time, they offer specific challenges in terms of data imputation. Here we compare statistical imputation approaches, using varying levels of environmental information, for five plant traits (leaf biomass to sapwood area ratio, leaf nitrogen content, maximum tree height, leaf mass per area and wood density) in a spatially explicit plant trait dataset of temperate and Mediterranean tree species (Ecological and Forest Inventory of Catalonia, IEFC, dataset for Catalonia, north-east Iberian Peninsula, 31 900 km2). We simulated gaps at different missingness levels (10-80 %) in a complete trait matrix, and we used overall trait means, species means, k nearest neighbours (kNN), ordinary and regression kriging, and multivariate imputation using chained equations (MICE) to impute missing trait values. We assessed these methods in terms of their accuracy and of their ability to preserve trait distributions, multi-trait correlation structure and bivariate trait relationships. The relatively good performance of mean and species mean imputations in terms of accuracy masked a poor representation of trait distributions and multivariate trait structure. Species identity improved MICE imputations for all traits, whereas forest structure and topography improved imputations for some traits. No method performed best consistently for the five studied traits, but, considering all traits and performance metrics, MICE informed by relevant ecological variables gave the best results. However, at higher missingness (> 30 %), species mean imputations and regression kriging tended to outperform MICE for some traits. MICE informed by relevant ecological variables allowed us to fill the gaps in the IEFC incomplete dataset (5495 plots) and quantify imputation uncertainty. Resulting spatial patterns of the studied traits in Catalan forests were broadly similar when using species means, regression kriging or the best-performing MICE application, but some important discrepancies were observed at the local level. Our results highlight the need to assess imputation quality beyond just imputation accuracy and show that including environmental information in statistical imputation approaches yields more plausible imputations in spatially explicit plant trait datasets. © 2018 Author(s).
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.
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.
Poyatos R., Aguade D., Galiano L., Mencuccini M., Martinez-Vilalta J. (2013) Drought-induced defoliation and long periods of near-zero gas exchange play a key role in accentuating metabolic decline of Scots pine. New Phytologist. 200: 388-401.EnlaceDoi: 10.1111/nph.12278
Summary: Drought-induced defoliation has recently been associated with the depletion of carbon reserves and increased mortality risk in Scots pine (Pinus sylvestris). We hypothesize that defoliated individuals are more sensitive to drought, implying that potentially higher gas exchange (per unit of leaf area) during wet periods may not compensate for their reduced photosynthetic area. We measured sap flow, needle water potentials and whole-tree hydraulic conductance to analyse the drought responses of co-occurring defoliated and nondefoliated Scots pines in northeast Spain during typical (2010) and extreme (2011) drought conditions. Defoliated Scots pines showed higher sap flow per unit leaf area during spring, but were more sensitive to summer drought, relative to nondefoliated pines. This pattern was associated with a steeper decline in soil-to-leaf hydraulic conductance with drought and an enhanced sensitivity of canopy conductance to soil water availability. Near-homeostasis in midday water potentials was observed across years and defoliation classes, with minimum values of -2.5 MPa. Enhanced sensitivity to drought and prolonged periods of near-zero gas exchange were consistent with low levels of carbohydrate reserves in defoliated trees. Our results support the critical links between defoliation, water and carbon availability, and their key roles in determining tree survival and recovery under drought. © 2013 New Phytologist Trust.
Sterck F.J., Martínez-Vilalta J., Mencuccini M., Cochard H., Gerrits P., Zweifel R., Herrero A., Korhonen J.F., Llorens P., Nikinmaa E., Nolè A., Poyatos R., Ripullone F., Sass-Klaassen U. (2012) Understanding trait interactions and their impacts on growth in Scots pine branches across Europe. Functional Ecology. 26: 541-549.EnlaceDoi: 10.1111/j.1365-2435.2012.01963.x
Plants exhibit a wide variety in traits at different organizational levels. Intraspecific and interspecific studies have potential to demonstrate functional relationships and trade-offs amongst traits, with potential consequences for growth. However, the distinction between the correlative and functional nature of trait covariation presents a challenge because traits interact in complex ways. We present an intraspecific study on Scots pine branches and use functional multi-trait concepts to organize and understand trait interactions and their impacts on growth. Branch-level traits were assessed for 97 branches from 12 Scots pine sites across Europe. To test alternative hypotheses on cause-effect relationships between anatomical traits, hydraulic traits and branch growth, we measured for each branch: the tracheid hydraulic diameter, double cell wall thickness, cell lumen span area, wood density, cavitation vulnerability, wood-specific hydraulic conductivity, the leaf area to sapwood area ratio and branch growth. We used mixed linear effect models and path models to show how anatomical traits determine hydraulic traits and, in turn, how those traits influence growth. Tracheid hydraulic diameter was the best predictor of cavitation vulnerability (R 2=0·09 explained by path model) and specific conductivity (R 2=0·19) amongst anatomical traits. Leaf area to sapwood area ratio had the strongest direct effect on branch growth (R 2=0·19) and was positively associated with the tracheid hydraulic diameter (R 2=0·22). A number of bivariate correlations between traits could be explained by these functional relationships amongst traits. The plasticity in tracheid hydraulic diameter (10.0-15.1μm) and leaf area to sapwood area ratio (600-6051cm 2cm -2) and the maintenance of a minimum leaf water potential (between -2 and -2·5MPa) appear to drive the anatomical and hydraulic traits of Scots pine across Europe. These properties are major drivers of the functional trait network underlying the growth variation amongst pine branches and thus possibly contribute to the ecological success of pines at a local and continental scale. © 2012 The Authors. Functional Ecology © 2012 British Ecological Society.
Martínez-Vilalta J., Cochard H., Mencuccini M., Sterck F., Herrero A., Korhonen J.F.J., Llorens P., Nikinmaa E., Nolè A., Poyatos R., Ripullone F., Sass-Klaassen U., Zweifel R. (2009) Hydraulic adjustment of Scots pine across Europe. New Phytologist. 184: 353-364.EnlaceDoi: 10.1111/j.1469-8137.2009.02954.x
Summary The variability of branch-level hydraulic properties was assessed across 12 Scots pine populations covering a wide range of environmental conditions, including some of the southernmost populations of the species. The aims were to relate this variability to differences in climate, and to study the potential tradeoffs between traits. Traits measured included wood density, radial growth, xylem anatomy, sapwood- and leaf-specific hydraulic conductivity (KS and KL), vulnerability to embolism, leaf-to-sapwood area ratio (AL : AS), needle carbon isotope discrimination (Δ13C) and nitrogen content, and specific leaf area. Between-population variability was high for most of the hydraulic traits studied, but it was directly associated with climate dryness (defined as a combination of atmospheric moisture demand and availability) only for A L : AS, KL and Δ13C. Shoot radial growth and AL : AS declined with stand development, which is consistent with a strategy to avoid exceedingly low water potentials as tree size increases. In addition, we did not find evidence at the intraspecific level of some associations between hydraulic traits that have been commonly reported across species. The adjustment of Scots pine's hydraulic system to local climatic conditions occurred primarily through modifications of AL : AS and direct stomatal control, whereas intraspecific variation in vulnerability to embolism and leaf physiology appears to be limited. © 2009 New Phytologist.
Poyatos R., Martínez-Vilalta J., Čermák J., Ceulemans R., Granier A., Irvine J., Köstner B., Lagergren F., Meiresonne L., Nadezhdina N., Zimmermann R., Llorens P., Mencuccini M. (2007) Plasticity in hydraulic architecture of Scots pine across Eurasia. Oecologia. 153: 245-259.EnlaceDoi: 10.1007/s00442-007-0740-0
Widespread tree species must show physiological and structural plasticity to deal with contrasting water balance conditions. To investigate these plasticity mechanisms, a meta-analysis of Pinus sylvestris L. sap flow and its response to environmental variables was conducted using datasets from across its whole geographical range. For each site, a Jarvis-type, multiplicative model was used to fit the relationship between sap flow and photosynthetically active radiation, vapour pressure deficit (D) and soil moisture deficit (SMD); and a logarithmic function was used to characterize the response of stomatal conductance (G s) to D. The fitted parameters of those models were regressed against climatic variables to study the acclimation of Scots pine to dry/warm conditions. The absolute value of sap flow and its sensitivity to D and SMD increased with the average summer evaporative demand. However, relative sensitivity of G s to D (m/G s,ref, where m is the slope and G s,ref is reference G s at D = 1 kPa) did not increase with evaporative demand across populations, and transpiration per unit leaf area at a given D increased accordingly in drier/warmer climates. This physiological plasticity was linked to the previously reported climate- and size-related structural acclimation of leaf to sapwood area ratios. G s,ref, and its absolute sensitivity to D (m), tended to decrease with age/height of the trees as previously reported for other pine species. It is unclear why Scots pines have higher transpiration rates at drier/warmer sites, at the expense of lower water-use efficiency. In any case, our results suggest that these structural adjustments may not be enough to prevent lower xylem tensions at the driest sites. © 2007 Springer-Verlag.
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