Tadeja Savi

Rooting depth, water relations and non-structural carbohydrate dynamics in three woody angiosperms differentially affected by an extreme summer drought

In 2012, an extreme summer drought induced species-specific die-back in woody species in Northeastern Italy. Quercus pubescens and Ostrya carpinifolia were heavily impacted, while Prunus mahaleb was largely unaffected. By comparing seasonal changes in isotopic composition of xylem sap, rainfall and deep soil samples, we show that P. mahaleb has a deeper root system than the other two species. This morphological trait allowed P  mahaleb to maintain higher water potential (Ψ), gas exchange rates and non-structural carbohydrates content (NSC) throughout the summer, when compared with the other species. More favourable water and carbon states allowed relatively stable maintenance of stem hydraulic conductivity (k) throughout the growing season. In contrast, in Quercus pubescens and Ostrya carpinifolia, decreasing Ψ and NSC were associated with significant hydraulic failure, with spring-to-summer k loss averaging 60%. Our data support the hypothesis that drought-induced tree decline is a complex phenomenon that cannot be modelled on the basis of single predictors of tree status like hydraulic efficiency, vulnerability and carbohydrate content. Our data highlight the role of rooting depth in seasonal progression of water status, gas exchange and NSC, with possible consequences for energy-demanding mechanisms involved in the maintenance of vascular integrity.

Diurnal changes in embolism rate in nine dry forest trees: relationships with species-specific xylem vulnerability, hydraulic strategy and wood traits

Recent studies have reported correlations between stem sapwood capacitance (C(wood)) and xylem vulnerability to embolism, but it is unclear how C(wood) relates to the eventual ability of plants to reverse embolism. We investigated possible functional links between embolism reversal efficiency, C(wood), wood density (WD), vulnerability to xylem embolism and hydraulic safety margins in nine woody species native to dry sclerophyllous forests with different degrees of iso versus anisohydry. Substantial inter-specific differences in terms of seasonal/diurnal changes of xylem and leaf water potential, maximum diurnal values of transpiration rate and xylem vulnerability to embolism formation were recorded. Significant diurnal changes in percentage loss of hydraulic conductivity (PLC) were recorded for five species. Significant correlations were recorded between diurnal PLC changes and P50 and P88 values (i.e., xylem pressure inducing 50 and 88% PLC, respectively) as well as between diurnal PLC changes and safety margins referenced to P50 and P88. WD was linearly correlated with minimum diurnal leaf water potential, diurnal PLC changes and wood capacitance across all species. In contrast, significant relationships between P50, safety margin values referenced to P50 and WD were recorded only for the isohydric species. Functional links between diurnal changes in PLC, hydraulic strategies and WD and C(wood) are discussed.

Drought-induced xylem cavitation and hydraulic deterioration: risk factors for urban trees under climate change?

Urban trees help towns to cope with climate warming by cooling both air and surfaces. The challenges imposed by the urban environment, with special reference to low water availability due to the presence of extensive pavements, result in high rates of mortality of street trees, that can be increased by climatic extremes. We investigated the water relations and xylem hydraulic safety/efficiency of Quercus ilex trees growing at urban sites with different percentages of surrounding impervious pavements. Seasonal changes of plant water potential and gas exchange, vulnerability to cavitation and embolism level, and morpho-anatomical traits were measured. We found patterns of increasing water stress and vulnerability to drought at increasing percentages of impervious pavement cover, with a consequent reduction in gas exchange rates, decreased safety margins toward embolism development, and increased vulnerability to cavitation, suggesting the occurrence of stress-induced hydraulic deterioration. The amount of impermeable surface and chronic exposure to water stress influence the site-specific risk of drought-induced dieback of urban trees under extreme drought. Besides providing directions for management of green spaces in towns, our data suggest that xylem hydraulics is key to a full understanding of the responses of urban trees to global change.

The contribution of vascular and extra-vascular water pathways to drought-induced decline of leaf hydraulic conductance.

Drought stress can impair leaf hydraulic conductance (Kleaf), but the relative contribution of changes in the efficiency of the vein xylem water pathway and in the mesophyll route outside the xylem in driving the decline of Kleaf is still debated. We report direct measurements of dehydration-induced changes in the hydraulic resistance (R=1/K) of whole leaf (Rleaf), as well as of the leaf xylem (Rx) and extra-vascular pathways (Rox) in four Angiosperm species. Rleaf, Rx, and Rox were measured using the vacuum chamber method (VCM). Rleaf values during progressive leaf dehydration were also validated with measurements performed using the rehydration kinetic method (RKM). We analysed correlations between changes in Rx or Rox and Rleaf, as well as between morpho-anatomical traits (including dehydration-induced leaf shrinkage), vulnerability to embolism, and leaf water relation parameters. Measurements revealed that the relative contribution of vascular and extra-vascular hydraulic properties in driving Kleaf decline during dehydration is species-specific. Whilst in two study species the progressive impairment of both vascular and extra-vascular pathways contributed to leaf hydraulic vulnerability, in the other two species the vascular pathway remained substantially unaltered during leaf dehydration, and Kleaf decline was apparently caused only by changes in the hydraulic properties of the extra-vascular compartment.

Green roofs for a drier world: effects of hydrogel amendment on substrate and plant water status.

Climate features of the Mediterranean area make plant survival over green roofs challenging, thus calling for research work to improve water holding capacities of green roof systems. We assessed the effects of polymer hydrogel amendment on the water holding capacity of a green roof substrate, as well as on water status and growth of Salvia officinalis. Plants were grown in green roof experimental modules containing 8 cm or 12 cm deep substrate (control) or substrate mixed with hydrogel at two different concentrations: 0.3 or 0.6%. Hydrogel significantly increased the substrates water content at saturation, as well as water available to vegetation. Plants grown in 8 cm deep substrate mixed with 0.6% of hydrogel showed the best performance in terms of water status and membrane integrity under drought stress, associated to the lowest above-ground biomass. Our results provide experimental evidence that polymer hydrogel amendments enhance water supply to vegetation at the establishment phase of a green roof. In particular, the water status of plants is most effectively improved when reduced substrate depths are used to limit the biomass accumulation during early growth stages. A significant loss of water holding capacity of substrate-hydrogel blends was observed after 5 months from establishment of the experimental modules. We suggest that cross-optimization of physical-chemical characteristics of hydrogels and green roof substrates is needed to improve long term effectiveness of polymer-hydrogel blends.

Plant performance on Mediterranean green roofs: interaction of species-specific hydraulic strategies and substrate water relations

Mediterranean native shrubs can be very useful for green roofs in hot and arid regions. Our data show that both Arbutus unedo L. and Salvia officinalis L. could be successfully utilized, although the choice of species should be based on the drought-resistant strategy relative to the desired technical performance of the green roof. Moreover, substrate selection was found to have a crucial role in the success of green roof installations in the Mediterranean area.

Species-specific reversal of stem xylem embolism after a prolonged drought correlates to endpoint concentration of soluble sugars

Recent reports on tree mortality associated with anomalous drought and heat have raised interest into processes underlying tree resistance/resilience to water stress. Hydraulic failure and carbon starvation have been proposed as main causes of tree decline, with recent theories treating water and carbon metabolism as interconnected processes. We subjected young plants of two native (Quercus pubescens [Qp] and Prunus mahaleb [Pm]) and two invasive (Robinia pseudoacacia [Rp] and Ailanthus altissima [Aa]) woody angiosperms to a prolonged drought leading to stomatal closure and xylem embolism, to induce carbon starvation and hydraulic failure. At the end of the treatment, plants were measured for embolism rates and NSC content, and re-irrigated to monitor recovery of xylem hydraulics. Data highlight different hydraulic strategies in native vs invasive species under water stress, and provide physiological explanations for species-specific impacts of recent severe droughts. Drought-sensitive species (Qp and Rp) suffered high embolism rates and were unable to completely refill xylem conduits upon restoration of water availability. Species that better survived recent droughts were able to limit embolism build-up (Pm) or efficiently restored hydraulic functionality after irrigation (Aa). Species-specific capacity to reverse xylem embolism correlated to stem-level concentration of soluble carbohydrates, but not to starch content.

Effects of prescribed burning on ecophysiological, anatomical and stem hydraulic properties in Pinus pinea L.

Prescribed burning (PB) is a widespread management technique for wildfire hazard abatement. Understanding PB effects on tree ecophysiology is key to defining burn prescriptions aimed at reducing fire hazard in Mediterranean pine plantations, such as Pinus pinea L. stands. We assessed physiological responses of adult P. pinea trees to PB using a combination of dendroecological, anatomical, hydraulic and isotopic analyses. Tree-ring widths, xylem cell wall thickness, lumen area, hydraulic diameter and tree-ring δ(13)C and δ(18)O were measured in trees on burned and control sites. Vulnerability curves were elaborated to assess tree hydraulic efficiency or safety. Despite the relatively intense thermal treatment (the residence time of temperatures above 50 °C at the stem surface ranged between 242 and 2239 s), burned trees did not suffer mechanical damage to stems, nor significant reduction in radial growth. Moreover, the PB did not affect xylem structure and tree hydraulics. No variations in (13)C-derived water use efficiency were recorded. This confirmed the high resistance of P. pinea to surface fire at the stem base. However, burned trees showed consistently lower δ(18)O values in the PB year, as a likely consequence of reduced competition for water and nutrients due to the understory burning, which increased both photosynthetic activity and stomatal conductance. Our multi-approach analysis offers new perspectives on post-fire survival strategies of P. pinea in an environment where fires are predicted to increase in frequency and severity during the 21st century.

Drought versus heat: What's the major constraint on Mediterranean green roof plants?

Green roofs are gaining momentum in the arid and semi-arid regions due to their multiple benefits as compared with conventional roofs. One of the most critical steps in green roof installation is the selection of drought and heat tolerant species that can thrive under extreme microclimate conditions. We monitored the water status, growth and survival of 11 drought-adapted shrub species grown on shallow green roof modules (10 and 13cm deep substrate) and analyzed traits enabling plants to cope with drought (symplastic and apoplastic resistance) and heat stress (root membrane stability). The physiological traits conferring efficiency/safety to the water transport system under severe drought influenced plant water status and represent good predictors of both plant water use and growth rates over green roofs. Moreover, our data suggest that high substrate temperature represents a stress factor affecting plant survival to a larger extent than drought per se. In fact, the major cause influencing seedling survival on shallow substrates was the species-specific root resistance to heat, a single and easy measurable trait that should be integrated into the methodological framework for screening and selection of suitable shrub species for roof greening in the Mediterranean.

Leaf hydraulic vulnerability protects stem functionality under drought stress in Salvia officinalis

Functional coordination between leaf and stem hydraulics has been proposed as a key trait of drought-resistant plants. A balanced water transport efficiency and safety of different plant organs might be of particular importance for plant survival in the Mediterranean climate. We monitored seasonal changes of leaf and stem water relations of Salvia officinalis L. in order to highlight strategies adopted by this species to survive in harsh environmental conditions. During summer drought, the water potential dropped below the turgor loss point thus reducing water loss by transpiration, whereas the photosynthetic efficiency remained relatively high. Leaves lost their water transport efficiency earlier than stems, although in both plant organs P50 (water potential inducing 50% loss of hydraulic conductivity) indicated surprisingly high vulnerability when compared with other drought-tolerant species. The fast recovery of leaf turgor upon restoration of soil water availability suggests that the reduction of leaf hydraulic conductance is not only a consequence of vein embolism, but cell shrinkage and consequent increase of resistance in the extra-xylem pathway may play an important role. We conclude that the drought tolerance of S. officinalis arises at least partly as a consequence of vulnerability segmentation.