Amplifying effects of recurrent drought on the dynamics of tree growth and water use in a subalpine forest.

Despite recent advances in our understanding of drought impacts on tree functioning, we lack knowledge about the dynamic responses of mature trees to recurrent drought stress. At a subalpine forest site, we assessed the effects of three years of recurrent experimental summer drought on tree growth and water relations of Larix decidua Mill. and Picea abies (L. Karst.), two common European conifers representative for contrasting water-use strategies. We combined dendrometer and xylem sap flow measurements with analyses of xylem anatomy and non-structural carbohydrates and their carbon-isotope composition. Recurrent drought increased the effects of soil moisture limitation on growth and xylogenesis, and to a lesser extent on xylem sap flow. P. abies showed stronger growth responses to recurrent drought, reduced starch concentrations in branches and increased water-use efficiency when compared to L. decidua. Despite comparatively larger maximum tree water deficits than in P. abies, xylem formation of L. decidua was less affected by drought, suggesting a stronger capacity of rehydration or lower cambial turgor thresholds for growth. Our study shows that recurrent drought progressively increases impacts on mature trees of both species, which suggests that in a future climate increasing drought frequency could impose strong legacies on carbon and water dynamics of treeline species.

Stem girdling indicates prioritized carbon allocation to the root system at the expense of radial stem growth in Norway spruce under drought conditions.

The early culmination of maximum radial growth (RG) in late spring has been found in several coniferous species in a dry inner Alpine environment. We hypothesized that an early decrease in RG is an adaptation to cope with drought stress, which might require an early switch of carbon (C) allocation to belowground organs. To test this hypothesis, we experimentally subjected six-year-old Norway spruce saplings (tree height: 1.35 m; n = 80 trees) to two levels of soil water availability (watered versus drought conditions) and manipulated tree C status by physically blocking phloem transport at three girdling dates (GD). The influence of C availability and drought on tree growth (radial and shoot growth; root biomass) in response to girdling was analyzed in both treatments. Non-structural carbohydrates (NSCs, soluble sugars and starch) were measured in the stem, root and current leader to evaluate changes in tree C status due to girdling. The main finding was a significant increase in RG of the girdled trees compared to the controls above the girdling zone (UZ). At all girdling dates the RG increase was significantly more intense in the drought-stressed compared with watered trees (c. 3.3 and 1.9-fold higher compared with controls in the drought-stressed and watered trees, respectively), most likely indicating that an early switch of C allocation to belowground occurs as an adaptation to maintain tree water status under drought conditions. Reactivation of the cambium after the cessation of its regular activity was detected in UZ in drought-stressed trees, while below the girdling zone no xylem formation was found and the NSC content was strikingly reduced. Irrespective of water availability, girdling before growth onset significantly reduced the progression of bud break (P < 0.05) and the length of the current leader shoot by -47% (P < 0.01) indicating a reduction in xylem hydraulic conductance, which was corroborated by significantly reduced xylem sap flow (P < 0.001). Based on our findings, we conclude that during the growing season drought stress prioritizes an early switch of C allocation to the root system as an adaptation to maintain adequate tree water status in drought-prone environments.

Environmental effects on stem water deficit in co-occurring conifers exposed to soil dryness.

We monitored dynamics of stem water deficit (ΔW) and needle water potential (Ψ) during two consecutive growing seasons (2011 and 2012) in a dry inner Alpine environment (750 m above sea level, Tyrol, Austria), where Pinus sylvestris, Picea abies and Larix decidua form mixed stands. ΔW was extracted from stem circumference variations, which were continuously recorded by electronic band dendrometers (six trees per species) and correlations with environmental variables were performed. Results revealed that (i) ΔW reached highest and lowest values in P. abies and L. decidua, respectively, while mean minimum water potential (Ψ(ea)) amounted to -3.0 MPa in L. decidua and -1.8 MPa in P. abies and P. sylvestris. (ii) ΔW and Ψ(ea) were significantly correlated in P. abies (r = 0.630; P = 0.038) and L. decidua (r = 0.646; P = 0.032). (iii) In all species, ΔW reached highest values in late summer and was most closely related to temperature (P < 0.001). Results indicate that all species were undergoing water limitations as measured by increasing ΔW throughout the growing season, whereby P. abies most strongly drew upon water reserves in the living tissues of the bark. Quite similar ΔW developed in drought-sensitive L. decidua and drought-tolerant P. sylvestris indicate that various water storage locations are depleted in species showing different strategies of water status regulation, i.e. anisohydric vs. isohydric behavior, respectively, and/or water uptake efficiency differs among these species. Close coupling of ΔW to temperature suggests that climate warming affects plant water status through its effect on atmospheric demand for moisture.

Transpiration and canopy conductance in an inner alpine Scots pine (Pinus sylvestris L.) forest.

Canopy transpiration (Ec) of a 150-year old Pinus sylvestris L. stand in an inner alpine dry valley, Tyrol, Austria was estimated throughout two growing seasons 2011 and 2012 by means of xylem sap flow measurements. Although there were prolonged periods of limited soil water availability Ec did not show a clear trend with respect to soil water availability and averaged 0.4 ± 0.19 mm day-1 under conditions of non-limiting soil water availability and 0.37 ± 0.17 mm day-1 when soil water availability was limited. This is because canopy conductance declined significantly with increasing evaporative demand and thus significantly reduced tree water loss. The growing season total of Ec was 74 mm and 88 mm in 2011 and 2012, respectively, which is significantly below the values estimated for other P. sylvestris forest ecosystems in Central Europe, and thus reflecting a strong adaptation to soil drought during periods of high evaporative.

Seasonal and Daily Xylem Radius Variations in Scots Pine Are Closely Linked to Environmental Factors Affecting Transpiration.

Seasonal and daily radius variations in the xylem (XRV) and inner bark (IBV) of mature Scots pine trees (Pinus sylvestris) were determined during April 2019-October 2021 at a drought-prone inner alpine site (c. 750 m asl; Tyrol, Austria) by applying point dendrometers. XRVs were also related to environmental factors to evaluate the drivers of XRV during the growing season. XRV records revealed that the xylem width (i) started to shrink around the onset of radial stem growth in April, (ii) consistently decreased by c. 50 µm at the time when air temperature (T) and vapor pressure deficit (VPD) reached their maximum in late June through mid-July, and (iii) recovered until November/December. Although in daily cycles of radius variations XRV preceded IBV by about two hours and the daily amplitude of XRV was about 1/10 that of IBV, XRV and IBV (seasonal trends removed) were closely linked (ρ = 0.755; p < 0.001), indicating tight hydraulic coupling between these tissues. Furthermore, the daily amplitude of XRV was linearly and closely related to daily maximum T (ρ = 0.802; p < 0.001), mean daily solar radiation (ρ = 0.809; p < 0.001), and non-linearly related to daily maximum VPD (R2= 0.837; p < 0.001), indicating that the xylem of Pinus sylvestris reacts like a transpiration-driven passive hydraulic system.

Climate Overrides the Influence of Microsite Conditions on Radial Growth of the Tall Multi-Stemmed Shrub Alnus alnobetula at Treeline.

Green alder (Alnus alnobetula), a tall multi-stemmed deciduous shrub, is widespread at high elevations in the Central European Alps. Its growth form frequently leads to asymmetric radial growth and anomalous growth ring patterns, making development of representative ring-width series a challenge. In order to assess the variability among radii of one shoot, among shoots belonging to one stock and among stocks, 60 stem discs were sampled at treeline on Mt. Patscherkofel (Tyrol, Austria). Annual increments were measured along 188 radii and analyzed in terms of their variability by applying dendrochronological techniques. Results revealed a high agreement in ring-width variation among radii of one shoot, among shoots of one stock and largely among stocks from different sites, confirming the pronounced limitation of radial stem growth by climate forcing at the alpine treeline. In contrast to this, a high variability in both absolute growth rates and long-term growth trends was found, which we attribute to different microsite conditions and disturbances. These factors also override climate control of radial growth under growth-limiting environmental conditions. Based on our findings we provide recommendations for the number of samples needed to carry out inter- and intra-annual studies of radial growth in this multi-stemmed clonal shrub.

Multivariate analysis of physiological parameters reveals a consistent O3 response pattern in leaves of adult European beech (Fagus sylvatica).

• Increasing atmospheric concentrations of phytotoxic ozone (O(3) ) can constrain growth and carbon sink strength of forest trees, potentially exacerbating global radiative forcing. Despite progress in the conceptual understanding of the impact of O(3) on plants, it is still difficult to detect response patterns at the leaf level. • Here, we employed principal component analysis (PCA) to analyse a database containing physiological leaf-level parameters of 60-yr-old Fagus sylvatica (European beech) trees. Data were collected over two climatically contrasting years under ambient and twice-ambient O(3) regimes in a free-air forest environment. • The first principal component (PC1) of the PCA was consistently responsive to O(3) and crown position within the trees over both years. Only a few of the original parameters showed an O(3) effect. PC1 was related to parameters indicative of oxidative stress signalling and changes in carbohydrate metabolism. PC1 correlated with cumulative O(3) uptake over preceding days. • PC1 represents an O(3) -responsive multivariate pattern detectable in the absence of consistently measurable O(3) effects on individual leaf-level parameters. An underlying effect of O(3) on physiological processes is indicated, providing experimental confirmation of theoretical O(3) response patterns suggested previously.

Respiratory fluxes in a Canary Islands pine forest.

We estimated component and whole-ecosystem CO(2) efflux (R(ECO)) in a Pinus canariensis Chr. Sm. ex DC stand in Tenerife, Canary Islands, an ecotone with strong seasonal changes in soil water availability. From November 2006 to February 2008, we measured foliage, stem and soil CO(2) efflux by chamber techniques. Site-specific CO(2) efflux models obtained from these chamber measurements were then combined with half-hourly measurements of canopy, stem and soil temperature as well as soil water potential, leaf and stem surface area data for scaling up component-specific CO(2) efflux to R(ECO). Integrated over an entire year, R(ECO) was 938 g of C m(-2) in 2007 and comprised the following component fluxes: 77% from soil, 11% from stems and 12% from foliage. Whole-ecosystem CO(2) efflux varied markedly throughout the year. During the cold and wet season, R(ECO) generally followed the seasonal trends in temperature, and during the warm and dry summer, however, R(ECO) was significantly reduced because of limited soil water availability in the main rooting horizon.

The water balance of grassland ecosystems in the Austrian Alps.

The altitudinal variation of precipitation, evapotranspiration and runoff was quantified at 16 different grassland sites between 580 and 2550 m a.s.l. in the Austrian Alps. Along this altitudinal transect annual evapotranspiration decreased from roughly 690 mm at low elevation sites to 210 - 220 mm at the upper limit of the alpine grassland belt. A detailed analysis of the data showed that the observed reduction in the annual sum of evapotranspiration could be mainly explained by the altitudinal decline of the length of the snow free period (i.e. the vegetation period). Daily mean sums of evapotranspiration showed no altitudinal trend and averaged 2.2 mm d-1 independent of elevation, although the leaf area index, growing season mean air temperature, and vapour pressure deficit declined with increasing altitude. As precipitation increased with elevation, evapotranspiration seems to be of secondary importance when compared to runoff. Inter-annual variability of evapotranspiration was fairly low across contrasting dry and wet years (coefficient of variation = 7 %), indicating that even during dry years water availability was not limiting evapotranspiration.

Long-term changes in tree-ring - climate relationships at Mt. Patscherkofel (Tyrol, Austria) since the mid 1980s.

Although growth limitation of trees at Alpine and high-latitude timberlines by prevailing summer temperature is well established, loss of thermal response of radial tree growth during last decades has repeatedly been addressed. We examined long-term variability of climate-growth relationships in ring width chronologies of Stone pine (Pinus cembra L.) by means of moving response functions (MRF). The study area is situated in the timberline ecotone (c. 2000 - 2200 m a.s.l.) on Mt. Patscherkofel (Tyrol, Austria). Five site chronologies were developed within the ecotone with constant sample depth (≥ 19 trees) throughout most of the time period analysed. MRF calculated for the period 1866-1999 and 1901-1999 for c. 200 and c. 100 yr old stands, respectively, revealed that mean July temperature is the major and long-term stable driving force of Pinus cembra radial growth within the timberline ecotone. However, since the mid 1980s, radial growth in timberline and tree line chronologies strikingly diverges from the July temperature trend. This is probably a result of extreme climate events (e.g. low winter precipitation, late frost) and/or increasing drought stress on cambial activity. The latter assumption is supported by a < 10 % increase in annual increments of c. 50 yr old trees at the timberline and at the tree line in 2003 compared to 2002, when extraordinary hot and dry conditions prevailed during summer. Furthermore, especially during the second half of the 20(th) century, influence of climate variables on radial growth show abrupt fluctuations, which might also be a consequence of climate warming on tree physiology.

Growing season water balance of an inner alpine Scots pine (Pinus sylvestris L.) forest.

We estimated components of the water cycle of a 150-year-old Pinus sylvestris forest in an inner Alpine dry valley of the Tyrol, Austria throughout five growing seasons. Forest canopy transpiration (TC) was measured by sap flow measurements scaled to the stand canopy level. Estimates of understory transpiration and forest floor evaporation (ETU) were derived from the soil water budget method, while interception (I) was modelled. Growing season cumulative evapotranspiration (ET = TC + ETU + I) varied between 256 and 322 mm or 51 to 79% of the growing season precipitation. The contribution of TC, ETU, and I to ET were 33, 40 and 27% respectively. Although these values of each layer (evapo)-transpiration are in good agreement with studies carried out in other European Scots pine forests, our estimated growing season total forest water use (Ttot = Tc + ETu) of 200-244 mm is at the lower end of values reported for coniferous forest ecosystems, and thus reflects an adaptation to the low shallow soil water availability. We conclude that Scots pine forests in inner alpine dry valleys are able to cope with high evaporative demand, even when shallow soil water availability is limited.

Nitrogen Addition and Understory Removal but Not Soil Warming Increased Radial Growth of Pinus cembra at Treeline in the Central Austrian Alps.

Beside low temperatures, limited tree growth at the alpine treeline may also be attributed to a lack of available soil nutrients and competition with understory vegetation. Although intra-annual stem growth of Pinus cembra has been studied intensively at the alpine treeline, the responses of radial growth to soil warming, soil fertilization, and below ground competition awaits clarification. In this study we quantified the effects of nitrogen (N) fertilization, soil warming, and understory removal on stem radial growth of P. cembra at treeline. Soil warming was achieved by roofing the forest floor with a transparent polyvinyl skin, while understory competition was prevented by shading the forest floor with a non-transparent foil around six trees each. Six trees received N- fertilization and six other trees served as controls. Stem growth was monitored with band dendrometers during the growing seasons 2012-2014. Our 3 years experiment showed that soil warming had no considerable effect on radial growth. Though understory removal through shading was accompanied by root-zone cooling, understory removal as well as N fertilization led to a significant increase in radial growth. Hardly affected was tree root biomass, while N-fertilization and understory removal significantly increased in 100-needle surface area and 100-needle dry mass, implying a higher amount of N stored in needles. Overall, our results demonstrate that beside low temperatures, tree growth at cold-climate boundaries may also be limited by root competition for nutrients between trees and understory vegetation. We conclude that tree understory interactions may also control treeline dynamics in a future changing environment.

Defense and avoidance of ozone under global change.

The level II approach of the critical loads concept adopted by the UNECE aims at a flux based evaluation and takes into account environmental factors governing stomatal conductance. These factors will probably be affected by global change. The flux concept predicts that a decrease in stomatal conductance would protect trees from air pollution effects by decreasing uptake. However, experimental evidence is inconclusive. Numerous results suggest that pollutants and factors subject to global change (drought, CO(2)) may interact and even exacerbate effects, probably because antioxidative defense systems are involved in both, defense against pollutant effects and protection from natural stress. An effective pollutant dose, which is weighted by physiological defense capacity, would better predict such effects. In this review paper we argue that the flux-based approach is imperfect, because global change effects may also modify the physiological susceptibility to ozone. Instead, a flux concept weighted by defense capacity should be tested.

Exemplifying whole-plant ozone uptake in adult forest trees of contrasting species and site conditions.

Whole-tree O3 uptake was exemplified for Picea abies, Fagus sylvatica and Larix decidua in stands at high and low altitude and contrasting water availability through sap flow measurement in tree trunks, intrinsically accounting for drought and boundary layer effects on O3 flux. O3 uptake of evergreen spruce per unit foliage area was enhanced by 100% at high relative to low elevation, whereas deciduous beech and larch showed similar uptake regardless of altitude. The responsiveness of the canopy conductance to water vapor and, as a consequence, O3 uptake to soil moisture and air humidity did not differ between species. Unifying findings at the whole-tree level will promote cause-effect based O3 risk assessment and modeling.

Quantification of ozone uptake at the stand level in a Pinus canariensis forest in Tenerife, Canary Islands: an approach based on sap flow measurements.

Ozone uptake was studied in a pine forest in Tenerife, Canary Islands, an ecotone with strong seasonal changes in climate. Ambient ozone concentration showed a pronounced seasonal course with high concentrations during the dry and warm period and low concentrations during the wet and cold season. Ozone uptake by contrast showed no clear seasonal trend. This is because canopy conductance significantly decreased with soil water availability and vapour pressure deficit. Mean daily ozone uptake averaged 1.9 nmol m(-2) s(-1) during the wet and cold season, and 1.5 nmol m(-2) s(-1) during the warm and dry period. The corresponding daily mean ambient ozone concentrations were 42 and 51 nl l(-1), respectively. Thus we conclude that in Mediterranean type forest ecosystems the flux based approach is more capable for risk assessment than an external, concentration based approach.

Stable Water Use Efficiency under Climate Change of Three Sympatric Conifer Species at the Alpine Treeline.

The ability of treeline associated conifers in the Central Alps to cope with recent climate warming and increasing CO2 concentration is still poorly understood. We determined tree ring stable carbon and oxygen isotope ratios of Pinus cembra, Picea abies, and Larix decidua trees from 1975 to 2010. Stable isotope ratios were compared with leaf level gas exchange measurements carried out in situ between 1979 and 2007. Results indicate that tree ring derived intrinsic water-use efficiency (iWUE) of P. cembra, P. abies and L. decidua remained constant during the last 36 years despite climate warming and rising atmospheric CO2. Temporal patterns in Δ(13)C and Δ(18)O mirrored leaf level gas exchange assessments, suggesting parallel increases of CO2-fixation and stomatal conductance of treeline conifer species. As at the study site soil water availability was not a limiting factor iWUE remained largely stable throughout the study period. The stability in iWUE was accompanied by an increase in basal area increment (BAI) suggesting that treeline trees benefit from both recent climate warming and CO2 fertilization. Finally, our results suggest that iWUE may not change species composition at treeline in the Austrian Alps due to similar ecophysiological responses to climatic changes of the three sympatric study species.

Global topics and novel approaches in the study of air pollution, climate change and forest ecosystems.

Research directions from the 27th conference for Specialists in Air Pollution and Climate Change Effects on Forest Ecosystems (2015) reflect knowledge advancements about (i) Mechanistic bases of tree responses to multiple climate and pollution stressors, in particular the interaction of ozone (O3) with nitrogen (N) deposition and drought; (ii) Linking genetic control with physiological whole-tree activity; (iii) Epigenetic responses to climate change and air pollution; (iv) Embedding individual tree performance into the multi-factorial stand-level interaction network; (v) Interactions of biogenic and anthropogenic volatile compounds (molecular, functional and ecological bases); (vi) Estimating the potential for carbon/pollution mitigation and cost effectiveness of urban and peri-urban forests; (vii) Selection of trees adapted to the urban environment; (viii) Trophic, competitive and host/parasite relationships under changing pollution and climate; (ix) Atmosphere-biosphere-pedosphere interactions as affected by anthropospheric changes; (x) Statistical analyses for epidemiological investigations; (xi) Use of monitoring for the validation of models; (xii) Holistic view for linking the climate, carbon, N and O3 modelling; (xiii) Inclusion of multiple environmental stresses (biotic and abiotic) in critical load determinations; (xiv) Ecological impacts of N deposition in the under-investigated areas; (xv) Empirical models for mechanistic effects at the local scale; (xvi) Broad-scale N and sulphur deposition input and their effects on forest ecosystem services; (xvii) Measurements of dry deposition of N; (xviii) Assessment of evapotranspiration; (xix) Remote sensing assessment of hydrological parameters; and (xx) Forest management for maximizing water provision and overall forest ecosystem services. Ground-level O3 is still the phytotoxic air pollutant of major concern to forest health. Specific issues about O3 are: (xxi) Developing dose-response relationships and stomatal O3 flux parameterizations for risk assessment, especially, in under-investigated regions; (xxii) Defining biologically based O3 standards for protection thresholds and critical levels; (xxiii) Use of free-air exposure facilities; (xxiv) Assessing O3 impacts on forest ecosystem services.

Increased water use efficiency does not prevent growth decline of Pinus canariensis in a semi-arid treeline ecotone in Tenerife, Canary Islands (Spain).

KEY MESSAGE: Intrinsic water-use efficiency of Pinus canariensis (Sweet ex Spreng.) growing at a semi-arid treeline has increased during the past 37 years. Tree-ring width by contrast has declined, likely caused by reduced stomatal conductance due to increasing aridity. CONTEXT: Rising atmospheric CO2 concentration (Ca ) has been related to tree growth enhancement accompanied by increasing intrinsic water-use-efficiency (iWUE). Nevertheless, the extent of rising Ca on long-term changes in iWUE and growth has remained poorly understood to date in Mediterranean treeline ecosystems. AIMS: This study aimed to examine radial growth and physiological responses of P. canariensis in relation to rising Ca and increasing aridity at treeline in Tenerife, Canary Islands, Spain. METHODS: We evaluated temporal changes in secondary growth (tree-ring width; TRW) and tree ring stable C isotope signature for assessing iWUE from 1975 through 2011. RESULTS: Precipitation was the main factor controlling secondary growth. Over the last 36 years P. canariensis showed a decline in TRW at enhanced iWUE, likely caused by reduced stomatal conductance due to increasing aridity. CONCLUSION: Our results indicate that increasing aridity has overridden the potential CO2 fertilization on tree growth of P. canariensis at its upper distribution limit.

Soil warming increased whole-tree water use of Pinus cembra at the treeline in the Central Tyrolean Alps.

This study quantified the effect of soil warming on sap flow density (Qs) of Pinus cembra L. at the treeline in the Central Tyrolean Alps. To enhance soil temperature we installed a transparent roof construction above the forest floor around six trees. Six other trees served as controls in the absence of any manipulation. Roofing enhanced growing season mean soil temperature by 1.6, 1.3 and 1.0 °C at 5, 10 and 20 cm soil depth, respectively, while soil water availability was not affected. Sap flow density (using Granier-type thermal dissipation probes) and environmental parameters were monitored throughout three growing seasons. During the first year of treatment, no warming effect was detected on Qs. However, soil warming caused Qs to increase significantly by 11 and 19% above levels in control trees during the second and third year, respectively. This effect appeared to result from warming-induced root production, a reduction in viscosity and perhaps an increase also in root hydraulic conductivity. Hardly affected were leaf-level net CO2 uptake rate and conductance for water vapour, so that water-use efficiency stayed unchanged as confirmed by needle δ(13)C analysis. We conclude that tree water loss will increase with soil warming, which may alter the water balance within the treeline ecotone of the Central Austrian Alps in a future warming environment.

Seasonal variation of soil respiration in a Pinus cembra forest at the upper timberline in the Central Austrian Alps.

Soil respiration (R) of a 95-year-old Pinus cembra L. forest at the alpine timberline was measured continuously from October 2001 to January 2003 with an automated multiplexing gas exchange system. There was significant spatial variability in soil respiration, and R at a soil temperature of 10 degrees C (R10) decreased by about 20% m(-1) with increasing distance from the trunk. Needle litter and fine root density also decreased. The spatially averaged annual soil CO2 efflux was 35 g C m(-2) year(-1) in 2002. About 70% of the temporal variation in soil respiration could be explained by variations in soil temperature, whereas the influence of soil water potential and thus soil water content was negligible because soil water availability was supra-optimal.