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Key message The lack of elevational changes in electrical resistivity in Pinus cembra trunks indicated consistent growth and hydraulics across elevations. Though, electrical resistivity tomograms exhibited pronounced temperature-driven seasonal changes. Abstract Alpine conifers growing at high elevation are exposed to low temperatures, which may limit xylogenesis and cause pronounced seasonal changes in tree hydraulics. Electrical resistivity (ER) tomography enables minimal invasive monitoring of stems in situ. We used this technique to analyze Pinus cembra trunks along a 400 m elevational gradient up to the timberline and over seasons. Furthermore, ER data of earlywood across tree rings were compared with the respective specific hydraulic conductivity (KS), measured on extracted wood cores. ER tomograms revealed pronounced changes over the year and a strong correlation between average ER (ERmean) and air and xylem temperatures. Surprisingly, no elevational changes in ERmean, earlywood ER or KS were observed. ER data corresponded to variation in earlywood KS, which decreased from the youngest (ca. 4–5 cm2s−1 MPa−1) to the oldest tree rings (0.63 ± 0.22 cm2s−1 MPa−1). The lack of changes in ER data and earlywood KS along the study transect indicated consistent growth patterns and no major changes in structural and functional hydraulic traits across elevation. The constant decrease in earlywood KS with tree ring age throughout all elevations highlights the hydraulic relevance of the outermost tree rings in P. cembra. Seasonal measurements demonstrated pronounced temperature effects on ER, and we thus recommend a detailed monitoring of trunk temperatures for ER tomography.

Xylem conductive capacity is a key determinant of plant hydraulic function and intimately linked to photosynthesis and productivity, but can be impeded by temporary or permanent conduit dysfunctions. Here we show that persistent xylem dysfunctions in unstressed plants are frequent in Alpine dwarf shrubs and occur in various but species-specific cross-sectional patterns. Combined synchrotron micro-computed tomography (micro-CT) imaging, xylem staining, and flow measurements in saturated samples of six widespread Ericaceae species evidence a high proportion (19%-50%) of hydraulically nonfunctional xylem areas in the absence of drought stress, with regular distribution of dysfunctions between or within growth rings. Dysfunctions were only partly reversible and reduced the specific hydraulic conductivity to 1.38 to 3.57 ×10-4 m2 s-1 MPa-1 . Decommission of inner growth rings was clearly related to stem age and a higher vulnerability to cavitation of older rings, while the high proportion of nonfunctional conduits in each annual ring needs further investigations. The lower the xylem fraction contributing to the transport function, the higher was the hydraulic efficiency of conducting xylem areas. Improved understanding of the functional lifespan of xylem elements and the prevalence and nature of dysfunctions is critical to correctly assess structure-function relationships and whole-plant hydraulic strategies.

Hydraulic conductance is recognized as a major determinant of gas exchange and productivity. However, whether this also applies to seedlings, a critically important stage for vegetation regeneration, has been largely unknown. We analyzed the hydraulic and stomatal conductance of leaves and shoots for 6-wk-old Acer pseudoplatanus seedlings emerging in different lowland and treeline habitats and under glasshouse conditions, respectively, as well as on 9-, 15- and 18-wk-old plants, and related findings to leaf and xylem anatomical traits. Treeline seedlings had higher leaf area-specific shoot hydraulic conductance (Kshoot-L ), and stomatal conductance (gs ), associated with wider xylem conduits, lower leaf area and higher stomatal density than lowland and glasshouse-grown plants. Across the first 18 wk of development, seedlings increased four-fold in absolute shoot hydraulic conductance (Kshoot ) and declined by half in Kshoot-L , with correlated shifts in xylem and leaf anatomy. Distal leaves had higher leaf hydraulic conductance (Kleaf ) and gs compared to basal leaves. Seedlings show strong variation across growth environments and ontogenetic shifts in hydraulic and anatomical parameters. Across growth sites, ontogenetic stages and leaf orders, gs was tightly correlated with Kshoot-L and Kleaf , balancing hydraulic supply with demand for the earliest stages of seedling establishment.

Dwarf shrubs are a key functional group of the high-elevation vegetation belt. Despite their ecological relevance and high sensitivity to environmental changes, the hydraulic adaptations and species-specific variations in drought tolerance within this growth form are mostly unknown. Here, we assess the xylem vulnerability to cavitation of important character species of the Alpine dwarf shrub heaths in Central Europe. Due to the high percentage of nonfunctional xylem areas in these species, vulnerability curves were gained by an innovative staining approach with safranin, validated by hydraulic and xylem anatomical measurements. The loss of 50% conducting xylem area appeared in the range of -1.78 to -2.91 MPa. Midday plant water potential during an intense summer drought remained above these critical thresholds but was correlated with cavitation resistance. No trade-off between hydraulic safety and efficiency was detected across analyzed species. We conclude that the characteristic clustered occurrence of species in the heterogeneous mountain terrain (mainly interpreted as a consequence of varying snow cover dynamics) is also reflected in species-specific adjustments in xylem hydraulics. The interspecific variety in vulnerability thresholds and plant water potentials during summer drought indicates different hydraulic strategies and adjustments in water relations between these co-occurring shrubs.

The seedling stage is the most susceptible one during a tree′s life. Water relations may be crucial for seedlings due to their small roots, limited water buffers and the effects of drought on water transport. Despite obvious relevance, studies on seedling xylem hydraulics are scarce as respective methodical approaches are limited. Micro‐CT scans of intact Acer pseudoplatanus and Fagus sylvatica seedlings dehydrated to different water potentials (Ψ) allowed the simultaneous observation of gas‐filled versus water‐filled conduits and the calculation of percentage loss of conductivity (PLC) in stems, roots and leaves (petioles or main veins). Additionally, anatomical analyses were performed and stem PLC measured with hydraulic techniques. In A. pseudoplatanus, petioles showed a higher Ψ at 50% PLC (Ψ50 −1.13MPa) than stems (−2.51 MPa) and roots (−1.78 MPa). The main leaf veins of F. sylvatica had similar Ψ50 values (−2.26 MPa) to stems (−2.74 MPa) and roots (−2.75 MPa). In both species, no difference between root and stems was observed. Hydraulic measurements on stems closely matched the micro‐CT based PLC calculations. Micro‐CT analyses indicated a species‐specific hydraulic architecture. Vulnerability segmentation, enabling a disconnection of the hydraulic pathway upon drought, was observed in A. pseudoplatanus but not in the especially shade‐tolerant F. sylvatica. Hydraulic patterns could partly be related to xylem anatomical traits.

High elevation spruce forests of the European Alps are frequently infected by the needle rust Chrysomyxa rhododendri, a pathogen causing remarkable defoliation, reduced tree growth and limited rejuvenation. Exact quantification of the disease severity on different spatial scales is crucial for monitoring, management and resistance breeding activities. Based on the distinct yellow discolouration of attacked needles, it was investigated whether image analysis of digital photographs can be used to quantify disease severity and to improve phenotyping compared to conventional assessment in terms of time, effort and application range. The developed protocol for preprocessing and analysis of digital RGB images enabled identification of disease symptoms and healthy needle areas on images obtained in ground surveys (total number of analysed images n = 62) and by the use of a semiprofessional quadcopter (n = 13). Obtained disease severities correlated linearly with results obtained by manual counting of healthy and diseased needles for all approaches, including images of individual branches with natural background (R 2 = 0.87) and with black background (R 2 = 0.95), juvenile plants (R 2 = 0.94), and top views and side views of entire tree crowns of adult trees (R 2 = 0.98 and 0.88, respectively). Results underline that a well‐defined signal related to needle bladder rust symptoms of Norway spruce can be extracted from images recorded by standard digital cameras and using drones. The presented protocol enables precise and time‐efficient quantification of disease symptoms caused by C. rhododendri and provides several advantages compared to conventional assessment by manual counting or visual estimations.

Key message Accumulation of phenolic needle metabolites in Norway spruce is regulated by many genes with small and additive effects and is correlated with the susceptibility against fungal attack. Abstract Norway spruce accumulates high foliar concentrations of secondary phenolic metabolites, with important functions for pathogen defence responses. However, the molecular genetic basis underlying the quantitative variation of phenolic compounds and their role in enhanced resistance of spruce to infection by needle bladder rust are unknown. To address these questions, a set of 1035 genome-wide single nucleotide polymorphisms (SNPs) was associated to the quantitative variation of four simple phenylpropanoids, eight stilbenes, nine flavonoids, six related arithmetic parameters and the susceptibility to infection by Chrysomyxa rhododendri in an unstructured natural population of Norway spruce. Thirty-one significant genetic associations for the flavonoids gallocatechin, kaempferol 3-glucoside and quercetin 3-glucoside and the stilbenes resveratrol, piceatannol, astringin and isorhapontin were discovered, explaining 22–59% of phenotypic variation, and indicating a regulation of phenolic accumulation by many genes with small and additive effects. The phenolics profile differed between trees with high and low susceptibility to the fungus, underlining the importance of phenolic compounds in the defence mechanisms of Norway spruce to C. rhododendri. Results highlight the utility of association studies in non-model tree species and may enable marker-assisted selection of Norway spruce adapted to severe pathogen attack. Electronic supplementary material The online version of this article (doi:10.1007/s11103-017-0589-5) contains supplementary material, which is available to authorized users.

Drought responses of mature trees are still poorly understood making it difficult to predict species distributions under a warmer climate. Using mature European beech (Fagus sylvatica L.), a widespread and economically important tree species in Europe, we aimed at developing an empirical stress-level scheme to describe its physiological response to drought. We analysed effects of decreasing soil and leaf water potential on soil water uptake, stem radius, native embolism, early defoliation and crown dieback with comprehensive measurements from overall nine hydrologically distinct beech stands across Switzerland, including records from the exceptional 2018 drought and the 2019/2020 post-drought period. Based on the observed responses to decreasing water potential we derived the following five stress levels: I (predawn leaf water potential-0.4 MPa): no detectable hydraulic limitations; II (-0.4 to -1.3): persistent stem shrinkage begins and growth ceases; III (-1.3 to -2.1): onset of native embolism and defoliation; IV (-2.1 to -2.8): onset of crown dieback; V (-2.8): transpiration ceases and crown dieback is20%. Our scheme provides, for the first time, quantitative thresholds regarding the physiological downregulation of mature European beech trees under drought and therefore synthesises relevant and fundamental information for process-based species distribution models. Moreover, our study revealed that European beech is drought vulnerable, because it still transpires considerably at high levels of embolism and because defoliation occurs rather as a result of embolism than preventing embolism. During the 2018 drought, an exposure to the stress levels III-V of only one month was long enough to trigger substantial crown dieback in beech trees on shallow soils. On deep soils with a high water holding capacity, in contrast, water reserves in deep soil layers prevented drought stress in beech trees. This emphasises the importance to include local data on soil water availability when predicting the future distribution of European beech.

Electrical resistivity (ER) tomography is a promising technique to minimally invasively study stems of living trees. It allows insights into xylem properties based on the cross-sectional distribution of ER that is governed by the wood’s electrical conductance. In this study, ER measurements were carried out on four forest tree species, Betula pendula, Fagus sylvatica, Picea abies and Pinus sylvestris, to demonstrate interspecific, intraspecific and within-tree variation of ER tomograms. Further, ER patterns were linked to xylem moisture content (MC), electrolyte content and density obtained from wood core analyses. The ER patterns of both coniferous species, P. abies and P. sylvestris, were found to be more homogenous and concentric compared with the complex tomograms of angiosperms. However, the ER range of coniferous trees showed considerable intraspecific variation. Measurements near ground level showed pronounced effects on ER tomograms, highlighting the importance of the chosen measurement height. A strong relation between ER and wood density was found in F. sylvatica while ER patterns of conifers were mainly influenced by MC. Results demonstrate a high species specificity of ER tomograms and of respective influencing xylem traits. They underline the importance of reference measurements for a correct interpretation of ER studies.

Common ivy ensures water supply of the crown by well-balanced within-plant adjustments of hydraulic efficiency and safety. Patterns correspond to variations in xylem anatomy and are similar to those of self-supporting trees., Lianas form long and flexible but disproportionately narrow stems, and thus require particular strategies to maintain the integrity of xylem water transport and ensure supply to large crown areas. The hydraulic architecture of lianas and the respective within-plant coordination of transport efficiency and safety, and the underlying anatomical variations in xylem, are largely unexplored. We analysed Hedera helix, a liana widespread in European temperate forests, with respect to hydraulic and xylem anatomical variations between the main stem and branches, between juvenile and adult life phases, and along the vertical axis. Main stems were significantly less embolism resistant but exhibited a higher hydraulic conductivity than branches. In branches, the cell turgor loss point of leaves decreased, while the embolism resistance and conductivity of xylem, as well as conduit diameters, increased with height. High water-transport capacities allow ivy to compensate for the small cross-section of stems, while the limited resistance to drought-induced xylem dysfunction of the main stem is probably linked to conservative stomatal regulation. Pronounced differences in xylem anatomy, hydraulic efficiency, and safety between the main stem and branches and along the vertical axis are surprisingly similar to those of self-supporting plants, and indicate that the coordination of carbon and water economies requires similar internal adjustments in tall plants.

Synchrotron X-ray computed micro-tomography (microCT) has emerged as a promising noninvasive technique for in vivo monitoring of xylem function, including embolism build-up under drought and hydraulic recovery following re-irrigation. Yet, the possible harmful effects of ionizing radiation on plant tissues have never been quantified. We specifically investigated the eventual damage suffered by stem living cells of three different species exposed to repeated microCT scans. Stem samples exposed to one, two or three scans were used to measure cell membrane and RNA integrity, and compared to controls never exposed to X-rays. Samples exposed to microCT scans suffered serious alterations to cell membranes, as revealed by marked increase in relative electrolyte leakage, and also underwent severe damage to RNA integrity. The negative effects of X-rays were apparent in all species tested, but the magnitude of damage and the minimum number of scans inducing negative effects were species-specific. Our data show that multiple microCT scans lead to disruption of fundamental cellular functions and processes. Hence, microCT investigation of phenomena that depend on physiological activity of living cells may produce erroneous results and lead to incorrect conclusions.

In alpine regions, tree hydraulics are limited by low temperatures that restrict xylem growth and induce winter frost drought and freezing stress. While several studies have dealt with functional limitations, data on elevational changes in functionally relevant xylem anatomical parameters are still scarce. In wood cores of Pinus cembra L. and Picea abies (L.) Karst. trunks, harvested along five elevational transects, xylem anatomical parameters (tracheid hydraulic diameter dh, wall reinforcement (t/b)2), pit dimensions (pit aperture Da, pit membrane Dm and torus Dt diameters) and respective functional indices (torus overlap O, margo flexibility) were measured. In both species, tracheid diameters decreased and (t/b)2 increased with increasing elevation, while pit dimensions and functional indices remained rather constant (P. cembra: Dt 10.3 ± 0.2 μm, O 0.477 ± 0.005; P. abies: Dt 9.30 ± 0.18 μm, O 0.492 ± 0.005). However, dh increased with tree height following a power trajectory with an exponent of 0.21, and also pit dimensions increased with tree height (exponents: Dm 0.18; Dt 0.14; Da 0.11). Observed elevational trends in xylem structures were predominantly determined by changes in tree size. Tree height-related changes in anatomical traits showed a remarkable robustness, regardless of the distributional ranges of study species. Despite increasing stress intensities towards the timberline, no adjustment in hydraulic safety at the pit level was observed.

Norway spruce trees in the subalpine forests of the European Alps are frequently attacked by the needle rust Chrysomyxa rhododendri. The obligate parasite undergoes a complex life cycle with a host shift between rhododendrons (Rhododendron sp.) and Norway spruce [Picea abies (L.) Karsten] and causes a yellowing and defoliation of the current-year needles in summer. Infected trees show several anatomical, morphological and physiological modifications, including a decrease in pigment content and net photosynthesis of infected needles, lower biomass production and reduced radial and height growth. The consequences are diminished timber yield and cripple growth. Because of repeated heavy infections in recent years, forest managers report increasing difficulties in both natural regeneration and afforestation at high elevation sites, where rhododendrons occur. This review gives a summary of the present knowledge about the effects of C. rhododendri infections on Norway spruce, including so far unpublished findings and with particular attention to the phenomena of resistant trees. Implications for subalpine forests and counter strategies are discussed.

The requirements of the water transport system of small herbaceous species differ considerably from those of woody species. Despite their ecological importance for many biomes, knowledge regarding herb hydraulics remains very limited. We compared key hydraulic features (vulnerability to drought-induced hydraulic decline, pressure-volume relations, onset of cellular damage, in situ variation of water potential, and stomatal conductance) of three Ranunculus species differing in their soil humidity preferences and ecological amplitude. All species were very vulnerable to water stress (50% reduction in whole-leaf hydraulic conductance [kleaf] at -0.2 to -0.8 MPa). In species with narrow ecological amplitude, the drought-exposed Ranunculus bulbosus was less vulnerable to desiccation (analyzed via loss of kleaf and turgor loss point) than the humid-habitat Ranunculus lanuginosus Accordingly, water stress-exposed plants from the broad-amplitude Ranunculus acris revealed tendencies toward lower vulnerability to water stress (e.g. osmotic potential at full turgor, cell damage, and stomatal closure) than conspecific plants from the humid site. We show that small herbs can adjust to their habitat conditions on interspecific and intraspecific levels in various hydraulic parameters. The coordination of hydraulic thresholds (50% and 88% loss of kleaf, turgor loss point, and minimum in situ water potential) enabled the study species to avoid hydraulic failure and damage to living cells. Reversible recovery of hydraulic conductance, desiccation-tolerant seeds, or rhizomes may allow them to prioritize toward a more efficient but vulnerable water transport system while avoiding the severe effects that water stress poses on woody species.

Vaccinium myrtillus and Vaccinium vitis‐idaea are two dwarf shrubs widespread in the European Alps. We studied the hydraulics of these species hypothesizing that (1) the hydraulic architecture of dwarf shrubs differs from trees, (2) hydraulic properties reflect the species' ecological amplitude and (3) hydraulic properties vary spatially and seasonally. Key hydraulic parameters (osmotic potential, turgor loss point, xylem hydraulic conductivity, vulnerability to drought‐induced embolism, stomata closure, drought‐induced cell damage and embolism repair) and related wood anatomical traits (conduit diameter and conduit wall reinforcement) were analyzed at four sites in Tyrol, Austria. Both species exhibited low hydraulic safety as well as low hydraulic efficiency. Fifty percentage embolism accumulated at −2.08 (V. myrtillus) and −1.97 MPa (V. vitis‐idaea), 88% stomata closure was at −2.19 and −2.35 MPa, respectively. After drought, both species showed embolism repair on re‐watering. Site‐specific variation within species was low, while seasonal changes in embolism resistance and turgor loss point were observed. Results indicate that studied Vaccinium species have a high risk for embolism formation. This is balanced by refilling capacities, which are probably based on the small growth height of dwarf shrubs. V. vitis‐idaea, which occurs on drier sites, showed more efficient repair and a lower turgor loss point than V. myrtillus.

Dwarf shrubs exhibit different requirements for a safe and efficient water supply compared to trees due their basitonic branching and low growth height. Though, only few studies dealt with the hydraulics of this growth form. Here we report key hydraulic parameters (vulnerability to drought-induced embolism, xylem hydraulic conductivity, cell osmotic potential, potential at turgor loss point) and related wood anatomical traits for Vaccinium gaultherioides, a wide-spread species in the European Alps. The results affirm the current knowledge, by indicating a relatively risky hydraulic strategy with low hydraulic safety compared to alpine trees and osmotic properties connected to the species’ soil humidity requirements.