Francesco Ripullone

Hydraulic adjustment of Scots pine across Europe.

* 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 (K(S) and K(L)), vulnerability to embolism, leaf-to-sapwood area ratio (A(L) : A(S)), needle carbon isotope discrimination (Delta13C) 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) : A(S), K(L) and Delta13C. Shoot radial growth and A(L) : A(S) 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 pines hydraulic system to local climatic conditions occurred primarily through modifications of A(L) : A(S) and direct stomatal control, whereas intraspecific variation in vulnerability to embolism and leaf physiology appears to be limited.

A multi-species synthesis of physiological mechanisms in drought-induced tree mortality

Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere–atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function.The mechanisms underlying drought-induced tree mortality are not fully resolved. Here, the authors show that, across multiple tree species, loss of xylem conductivity above 60% is associated with mortality, while carbon starvation is not universal.

Environmental Effects on Oxygen Isotope Enrichment of Leaf Water in Cotton Leaves

The oxygen isotope enrichment of bulk leaf water (Δb) was measured in cotton (Gossypium hirsutum) leaves to test the Craig-Gordon and Farquhar-Gan models under different environmental conditions. Δb increased with increasing leaf-to-air vapor pressure difference (VPd) as an overall result of the responses to the ratio of ambient to intercellular vapor pressures (ea/ei) and to stomatal conductance (gs). The oxygen isotope enrichment of lamina water relative to source water \batchmode \documentclass[fleqn,10pt,legalpaper]{article} \usepackage{amssymb} \usepackage{amsfonts} \usepackage{amsmath} \pagestyle{empty} \begin{document} \(({\bar{{\Delta}}}_{1}),\) \end{document} which increased with increasing VPd, was estimated by mass balance between less enriched water in primary veins and enriched water in the leaf. The Craig-Gordon model overestimated Δb (and \batchmode \documentclass[fleqn,10pt,legalpaper]{article} \usepackage{amssymb} \usepackage{amsfonts} \usepackage{amsmath} \pagestyle{empty} \begin{document} \({\bar{{\Delta}}}_{1}),\) \end{document} as expected. Such discrepancies increased with increase in transpiration rate (E), supporting the Farquhar-Gan model, which gave reasonable predictions of Δb and \batchmode \documentclass[fleqn,10pt,legalpaper]{article} \usepackage{amssymb} \usepackage{amsfonts} \usepackage{amsmath} \pagestyle{empty} \begin{document} \({\bar{{\Delta}}}_{1}\) \end{document} with an L of 7.9 mm, much less than the total radial effective length Lr of 43 mm. The fitted values of L for \batchmode \documentclass[fleqn,10pt,legalpaper]{article} \usepackage{amssymb} \usepackage{amsfonts} \usepackage{amsmath} \pagestyle{empty} \begin{document} \({\bar{{\Delta}}}_{1}\) \end{document} of individual leaves showed little dependence on VPd and temperature, supporting the assumption that the Farquhar-Gan formulation is relevant and useful in describing leaf water isotopic enrichment.

Assessing the effects of nitrogen deposition and climate on carbon isotope discrimination and intrinsic water-use efficiency of angiosperm and conifer trees under rising CO2 conditions.

The objective of this study is to globally assess the effects of atmospheric nitrogen deposition and climate, associated with rising levels of atmospheric CO2 , on the variability of carbon isotope discrimination (Δ(13) C), and intrinsic water-use efficiency (iWUE) of angiosperm and conifer tree species. Eighty-nine long-term isotope tree-ring chronologies, representing 23 conifer and 13 angiosperm species for 53 sites worldwide, were extracted from the literature, and used to obtain long-term time series of Δ(13) C and iWUE. Δ(13) C and iWUE were related to the increasing concentration of atmospheric CO2 over the industrial period (1850-2000) and to the variation of simulated atmospheric nitrogen deposition and climatic variables over the period 1950-2000. We applied generalized additive models and linear mixed-effects models to predict the effects of climatic variables and nitrogen deposition on Δ(13) C and iWUE. Results showed a declining Δ(13) C trend in the angiosperm and conifer species over the industrial period and a 16.1% increase of iWUE between 1850 and 2000, with no evidence that the increased rate was reduced at higher ambient CO2 values. The temporal variation in Δ(13) C supported the hypothesis of an active plant mechanism that maintains a constant ratio between intercellular and ambient CO2 concentrations. We defined linear mixed-effects models that were effective to describe the variation of Δ(13) C and iWUE as a function of a set of environmental predictors, alternatively including annual rate (Nrate ) and long-term cumulative (Ncum ) nitrogen deposition. No single climatic or atmospheric variable had a clearly predominant effect, however, Δ(13) C and iWUE showed complex dependent interactions between different covariates. A significant association of Nrate with iWUE and Δ(13) C was observed in conifers and in the angiosperms, and Ncum was the only independent term with a significant positive association with iWUE, although a multi-factorial control was evident in conifers.

Physiological and structural changes in response to altered precipitation regimes in a Mediterranean macchia ecosystem

Significant decrease in precipitation up to 15–20% has been observed in the Mediterranean area in the last two decades as a consequence of climate change. To simulate an analogous scenario, the precipitation regime was altered in replicated experimental plots in a Mediterranean macchia dominated by Arbutus unedo L. species. Two different levels of soil water content (SWC) were obtained during the summer: a mean value of 7% was obtained in water-depleted (D) plots by a partial (−20%) rain exclusion treatment using rain gutters; while a mean value of 14% in SWC was obtained in watered (W) plots supplying water by a sprinkler net. The physiological and structural changes were investigated over the course of two consecutive years by measurement of water potential, gas exchange leaf carbon isotopes, leaf pigments and growth. Apart from short-term responses, mainly related to the elastic response of stomatal conductance to soil water, a more long-lasting and significant acclimation to water availability was observed as a result of the increase in hydraulic resistance in the soil–plant continuum, which persisted even after the return to full water availability during the fall and winter. This response involved the permanent down-regulation of stomatal conductance and photosynthesis, accumulation of photo-protective pigments, as well as a reduction in shoot growth, leaf area index and an increase in shoot-bearing flowers in D plots. This acclimation response prevented the onset of any run-away damage thereby reducing the forest vulnerability to drought. Furthermore, the imposed drought induced a slight increase or no change in intrinsic water-use efficiency (WUEint), as a result of the parallel increase in stomatal and non-stomatal limitations; conversely integrated WUE (i.e., estimated from leaf carbon isotopes) was not affected by drought.

A multi-proxy assessment of dieback causes in a Mediterranean oak species

Drought stress causes forest dieback that is often explained by two interrelated mechanisms, namely hydraulic failure and carbon starvation. However, it is still unclear which functional and structural alterations, related to these mechanisms, predispose to dieback. Here we apply a multi-proxy approach for the characterization of tree structure (radial growth, wood anatomy) and functioning (δ13C, δ18O and non-structural carbohydrates (NSCs)) in tree rings before and after drought-induced dieback. We aim to discriminate which is the main mechanism and to assess which variables can act as early-warning proxies of drought-triggered damage. The study was tailored in southern Italy in two forests (i.e., San Paolo (SP) and Oriolo (OR)) where declining and non-declining trees of a ring-porous tree species (Quercus frainetto Ten.) showing anisohydric behavior coexist. Both stands showed growth decline in response to warm and dry spring conditions, although the onset of dieback was shifted between them (2002 in SP and 2009 in OR). Declining trees displayed a sharp growth drop after this onset with reductions of 49% and 44% at SP and OR sites, respectively. Further, contrary to what we expected, declining trees showed a lower intrinsic water-use efficiency compared with non-declining trees after the dieback onset (with reductions of 9.7% and 5.6% at sites SP and OR, respectively), due to enhanced water loss through transpiration, as indicated by the lower δ18O values. This was more noticeable at the most drought-affected SP stand. Sapwood NSCs did not differ between declining and non-declining trees, indicating no carbon starvation in affected trees. Thus, the characterized structural and functional alterations partially support the hydraulic failure mechanism of dieback. Finally, we show that growth data are reliable early-warning proxies of drought-triggered dieback.

Anthropogenic NOx emissions alter the intrinsic water-use efficiency (WUEi) for Quercus cerris stands under Mediterranean climate conditions.

We investigated the effect of N deposition (Ndep) on intrinsic water-use efficiency (WUEi), the ratio of photosynthesis (A) to stomatal conductance (gs), for two Quercus cerris stands at different distances to an oil refinery in Southern Italy. We used delta13C in tree rings for assessing changes in WUEi; while the influence of climate and NOx emission was explored through delta18O and delta15N, respectively. Differences in WUEi between the two sites were significant, with trees exposed to different degrees of NOx emissions showing an abrupt increase with the onset of pollution. Assuming similar gs at the two sites, as inferred through delta18O, the higher N availability at the polluted site caused the shift of the A/gs ratio in favour of A. Overall, our result suggests that an increase of Ndep may enhance tree WUE under a scenario of reduction of precipitation predicted for Mediterranean area.

Size Matters a Lot: Drought-Affected Italian Oaks Are Smaller and Show Lower Growth Prior to Tree Death

Hydraulic theory suggests that tall trees are at greater risk of drought-triggered death caused by hydraulic failure than small trees. In addition the drop in growth, observed in several tree species prior to death, is often interpreted as an early-warning signal of impending death. We test these hypotheses by comparing size, growth, and wood-anatomy patterns of living and now-dead trees in two Italian oak forests showing recent mortality episodes. The mortality probability of trees is modeled as a function of recent growth and tree size. Drift-diffusion-jump (DDJ) metrics are used to detect early-warning signals. We found that the tallest trees of the anisohydric Italian oak better survived drought contrary to what was predicted by the theory. Dead trees were characterized by a lower height and radial-growth trend than living trees in both study sites. The growth reduction of now-dead trees started about 10 years prior to their death and after two severe spring droughts during the early 2000s. This critical transition in growth was detected by DDJ metrics in the most affected site. Dead trees were also more sensitive to drought stress in this site indicating different susceptibility to water shortage between trees. Dead trees did not form earlywood vessels with smaller lumen diameter than surviving trees but tended to form wider latewood vessels with a higher percentage of vessel area. Since living and dead trees showed similar competition we did not expect that moderate thinning and a reduction in tree density would increase the short-term survival probability of trees.

Apical dominance ratio as an indicator of the growth conditions favouring Abies alba natural regeneration under Mediterranean environment

As reported in several studies, the presence of Abies alba Mill (silver fir) has been declining throughout its natural range over a large part of mountainous belt in the Mediterranean area. In such a context, regeneration establishment depends mainly on the occurrence of a suitable combination of water and light availability. Thus, before planning any forest management practice, it is essential to investigate on the optimal microclimate conditions influencing the success of natural regeneration of silver fir. To this aim, changes in growth and photosynthesis together with C, N and O isotope compositions have been investigated on silver fir naturally recruited saplings, growing in mixed stands with Fagus sylvatica on Apennine mountains (southern Italy). The apical dominance ratio (ADR, ratio between apical shoot length and length of first whorl lateral twigs) has been used as an indicator for microclimate conditions in which saplings grow. Based on the range of ADR values (i.e. from 0.10 to 1.30), saplings were distributed in four classes. As expected, increases in height, root collar diameter and radial growth correspond to enhancing ADR values, gaining the optimal conditions in class IV. This latter also displayed the best performance in terms of maximum CO2 assimilation at saturating light (Amax) and water-use efficiency as assessed by carbon isotope discrimination analysis. Conversely, class I and II seem to display the highest performance in terms of CO2 respiration rate (Rd) and absolute water loss saving as assessed by the application of oxygen isotopes. We conclude that, in relatively mild Mediterranean areas, forest managers should promote silvicultural treatments favouring light conditions and migration of saplings towards class IV of ADR. This class represents the optimal microclimate for regeneration establishment of silver fir.

It's a long way to the top: Plant species diversity in the transition from managed to old-growth forests

Questions Do vascular plant species richness and β-diversity differ between managed and structurally complex unmanaged stands? To what extent do species richness and β-diversity relate to forest structural attributes and heterogeneity?. Location Five National Parks in central and southern Italy. Methods We sampled vascular plant species composition and forest structural attributes in eight unmanaged temperate mesic forest stands dominated or co-dominated by beech, and in eight comparison stands managed as high forests with similar environmental features. We compared plant species richness, composition and β-diversity, across pairs of stands (unmanaged vs. managed) using Generalized Linear Mixed (effect) Models (GLMMs). β-diversity was quantified both at the scale of each pair of stands using plot-to-plot dissimilarity matrices (species turnover), and across the whole dataset, considering the distance in the multivariate species space of individual plots from the centroid of the plots within the same stand (compositional heterogeneity). We modelled the relationship between species diversity (richness and β-diversity) and forest structural heterogeneity and individual structural variables using GLMMs and Multiple Regression on Distance Matrices. Results Species composition differed significantly between managed and unmanaged stands, but not richness and β-diversity. We found weak evidence that plant species richness increased with increasing levels of structural heterogeneity and canopy diversification. At the scale of individual stands, species turnover was explained by different variables in distinct stands, with variables related to deadwood quantity and quality being selected most often. Conversely, we did not find support to the hypothesis that compositional heterogeneity varies as a function of forest structural characteristics at the scale of the whole dataset. Conclusions Structurally complex unmanaged stands have a distinct herb-layer species composition from that of mature stands in similar environmental conditions; nevertheless, we did not find significantly higher levels of vascular plant species richness and β-diversity in unmanaged stands. β-diversity was related to patterns of deadwood accumulation, while for species richness the evidence that it increases with increasing levels of canopy diversification was weak. These results suggest that emulating natural disturbance, and favoring deadwood accumulation and canopy diversification may benefit some, but not all facets of plant species diversity in Apennine beech forests. This article is protected by copyright. All rights reserved.