High preseason temperature variability drives convergence of xylem phenology in the Northern Hemisphere conifers.

Wood growth is key to understanding the feedback of forest ecosystems to the ongoing climate warming. An increase in spatial synchrony (i.e., coincident changes in distant populations) of spring phenology is one of the most prominent climate responses of forest trees. However, whether temperature variability contributes to an increase in the spatial synchrony of spring phenology and its underlying mechanisms remains largely unknown. Here, we analyzed an extensive dataset of xylem phenology observations of 20 conifer species from 75 sites over the Northern Hemisphere. Along the gradient of increase in temperature variability in the 75 sites, we observed a convergence in the onset of cell enlargement roughly toward the 5th of June, with a convergence in the onset of cell wall thickening toward the summer solstice. The increase in rainfall since the 5th of June is favorable for cell division and expansion, and as the most hours of sunlight are received around the summer solstice, it allows the optimization of carbon assimilation for cell wall thickening. Hence, the convergences can be considered as the result of matching xylem phenological activities to favorable conditions in regions with high temperature variability. Yet, forest trees relying on such consistent seasonal cues for xylem growth could constrain their ability to respond to climate warming, with consequences for the potential growing season length and, ultimately, forest productivity and survival in the future.

Incorporating high-resolution climate, remote sensing and topographic data to map annual forest growth in central and eastern Europe.

To enhance our understanding of forest carbon sequestration, climate change mitigation and drought impact on forest ecosystems, the availability of high-resolution annual forest growth maps based on tree-ring width (TRW) would provide a significant advancement to the field. Site-specific characteristics, which can be approximated by high-resolution Earth observation by satellites (EOS), emerge as crucial drivers of forest growth, influencing how climate translates into tree growth. EOS provides information on surface reflectance related to forest characteristics and thus can potentially improve the accuracy of forest growth models based on TRW. Through the modelling of TRW using EOS, climate and topography data, we showed that species-specific models can explain up to 52 % of model variance (Quercus petraea), while combining different species results in relatively poor model performance (R2 = 13 %). The integration of EOS into models based solely on climate and elevation data improved the explained variance by 6 % on average. Leveraging these insights, we successfully generated a map of annual TRW for the year 2021. We employed the area of applicability (AOA) approach to delineate the range in which our models are deemed valid. The calculated AOA for the established forest-type models was 73 % of the study region, indicating robust spatial applicability. Notably, unreliable predictions predominantly occurred in the climate margins of our dataset. In conclusion, our large-scale assessment underscores the efficacy of combining climate, EOS and topographic data to develop robust models for mapping annual TRW. This research not only fills a critical void in the current understanding of forest growth dynamics but also highlights the potential of integrated data sources for comprehensive ecosystem assessments.

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.

A critical thermal transition driving spring phenology of Northern Hemisphere conifers.

Despite growing interest in predicting plant phenological shifts, advanced spring phenology by global climate change remains debated. Evidence documenting either small or large advancement of spring phenology to rising temperature over the spatio-temporal scales implies a potential existence of a thermal threshold in the responses of forests to global warming. We collected a unique data set of xylem cell-wall-thickening onset dates in 20 coniferous species covering a broad mean annual temperature (MAT) gradient (-3.05 to 22.9°C) across the Northern Hemisphere (latitudes 23°-66° N). Along the MAT gradient, we identified a threshold temperature (using segmented regression) of 4.9 ± 1.1°C, above which the response of xylem phenology to rising temperatures significantly decline. This threshold separates the Northern Hemisphere conifers into cold and warm thermal niches, with MAT and spring forcing being the primary drivers for the onset dates (estimated by linear and Bayesian mixed-effect models), respectively. The identified thermal threshold should be integrated into the Earth-System-Models for a better understanding of spring phenology in response to global warming and an improved prediction of global climate-carbon feedbacks.

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.

Triggering Bimodal Radial Stem Growth in Pinus sylvestris at a Drought-Prone Site by Manipulating Stem Carbon Availability.

A bimodal radial growth (RG) pattern, i.e., growth peaks in spring and autumn, was repeatedly found in trees in the Mediterranean regions, where summer drought causes reduction or cessation of cambial activity. In a dry inner Alpine valley of the Eastern Alps (Tyrol, Austria, 750 m asl), Pinus sylvestris shows unimodal RG with onset and cessation of cambial activity in early April and late June, respectively. A resumption of cambial activity after intense summer rainfall was not observed in this region. In a field experiment, we tested the hypothesis that early cessation of cambial activity at this drought-prone site is an adaptation to limited water availability leading to an early and irreversible switch of carbon (C) allocation to belowground. To accomplish this, the C status of young P. sylvestris trees was manipulated by physical blockage of phloem transport (girdling) 6 weeks after cessation of cambial cell division. Influence of manipulated C availability on RG was recorded by stem dendrometers, which were mounted above the girdling zone. In response to blockage of phloem flow, resumption of cambial activity was detected above girdling after about 2 weeks. Although the experimentally induced second growth surge lasted for the same period as in spring (c. 2 months), the increment was more than twice as large due to doubling of daily maximum RG rate. After girdling, wood anatomical traits above girdling no longer showed any significant differences between earlywood and latewood tracheids indicating pronounced effects of C availability on cell differentiation. Below girdling, no reactivation of cambial activity occurred, but cell wall thickness of last formed latewood cell was reduced due to lack of C supply after girdling. Intense RG resumption after girdling indicates that cessation of cambial activity can be reversed by manipulating C status of the stem. Hence, our girdling study yielded strong support for the hypothesis that belowground organs exert high C sink strengths on the drought-prone study site. Furthermore, this work highlights the need of in-depth experimental studies in order to understand the interactions between endogenous and exogenous factors on cambial activity and xylem cell differentiation more clearly.

Tracheid and Pit Dimensions Hardly Vary in the Xylem of Pinus sylvestris Under Contrasting Growing Conditions.

Maintaining sufficient water transport via the xylem is crucial for tree survival under variable environmental conditions. Both efficiency and safety of the water transport are based on the anatomical structure of conduits and their connections, the pits. Yet, the plasticity of the xylem anatomy, particularly that of the pit structures, remains unclear. Also, trees adjust conduit dimensions to the water transport distance (i.e., tree size), but knowledge on respective adjustments in pit dimensions is scarce. We compared tracheid traits [mean tracheid diameter d, mean hydraulic diameter d h , cell wall reinforcement (t/b)2], pit dimensions (diameters of pit aperture D a , torus D t , margo D m , and pit border D p ), and pit functional properties (margo flexibility F, absolute overlap O a , torus overlap O, and valve effect V ef ) of two Scots pine (Pinus sylvestris L.) stands of similar tree heights but contrasting growth rates. Furthermore, we analyzed the trends of these xylem anatomical parameters across tree rings. Tracheid traits and pit dimensions were similar on both sites, whereas O a , O, and F were higher at the site with a lower growth rate. On the lower growth rate site, d h and pit dimensions increased across tree rings from pith to bark, and in trees from both sites, d h scaled with pit dimensions. Adjusted pit functional properties indicate slightly higher hydraulic safety in trees with a lower growth rate, although a lack of major differences in measured traits indicated overall low plasticity of the tracheid and pit architecture. Mean hydraulic diameter and pit dimension are well coordinated to increase the hydraulic efficiency toward the outer tree rings and thus with increasing tree height. Our results contribute to a better understanding of tree hydraulics under variable environmental conditions.

Photoperiod and temperature as dominant environmental drivers triggering secondary growth resumption in Northern Hemisphere conifers.

Wood formation consumes around 15% of the anthropogenic CO2 emissions per year and plays a critical role in long-term sequestration of carbon on Earth. However, the exogenous factors driving wood formation onset and the underlying cellular mechanisms are still poorly understood and quantified, and this hampers an effective assessment of terrestrial forest productivity and carbon budget under global warming. Here, we used an extensive collection of unique datasets of weekly xylem tissue formation (wood formation) from 21 coniferous species across the Northern Hemisphere (latitudes 23 to 67°N) to present a quantitative demonstration that the onset of wood formation in Northern Hemisphere conifers is primarily driven by photoperiod and mean annual temperature (MAT), and only secondarily by spring forcing, winter chilling, and moisture availability. Photoperiod interacts with MAT and plays the dominant role in regulating the onset of secondary meristem growth, contrary to its as-yet-unquantified role in affecting the springtime phenology of primary meristems. The unique relationships between exogenous factors and wood formation could help to predict how forest ecosystems respond and adapt to climate warming and could provide a better understanding of the feedback occurring between vegetation and climate that is mediated by phenology. Our study quantifies the role of major environmental drivers for incorporation into state-of-the-art Earth system models (ESMs), thereby providing an improved assessment of long-term and high-resolution observations of biogeochemical cycles across terrestrial biomes.

Hygroscopic properties of thin dead outer bark layers strongly influence stem diameter variations on short and long time scales in Scots pine (Pinus sylvestris L.).

Time series of stem diameter variations (SDVs) recorded by dendrometers are composed of two components: (i) irreversible radial stem growth and (ii) reversible stem shrinking and swelling caused by dynamics in water storage in elastic tissues outside the cambium. However, SDVs measured over dead outer bark (periderm) could also be affected by absorption and evaporation of water from remaining dead bark layers after smoothing the stem surface to properly mount dendrometers. Therefore, the focus of this study was to determine the influence of hygroscopicity of a thin dead outer bark layer on the reversible component of dendrometer records of Scots pine (Pinus sylvestris) under field conditions. To accomplish this, SDVs deduced from dendrometers mounted over dead outer bark were compared among living and dead saplings and mature trees. Results revealed that dead trees showed high synchronicity in reversible daily SDVs compared to living trees throughout several growing seasons (mean Pearson correlation coefficient (r) = 0.844 among saplings and r = 0.902 among mature trees, respectively; P<0.001). Furthermore, diurnal and long-term SDVs closely followed changes in relative air humidity (RH) in living and dead trees. A multiple linear regression analysis of environmental influence on SDVs in dead and living trees revealed that the most important predictor of daily SDVs was RH (relative importance 64 %). Hence, results indicate that dendrometers mounted over dead outer bark with a thickness of <4 mm record hygroscopic shrinking and swelling of the bark tissue, which can amplify fluctuations in whole-tree water status. To conclude, hygroscopic processes must be taken into account when extracting intra-annual radial growth, determining environmental drivers of SDVs, and evaluating changes in tree water status from SDVs recorded by dendrometers, which were mounted over even thin dead outer bark layers.

Low growth resilience to drought is related to future mortality risk in trees.

Severe droughts have the potential to reduce forest productivity and trigger tree mortality. Most trees face several drought events during their life and therefore resilience to dry conditions may be crucial to long-term survival. We assessed how growth resilience to severe droughts, including its components resistance and recovery, is related to the ability to survive future droughts by using a tree-ring database of surviving and now-dead trees from 118 sites (22 species, >3,500 trees). We found that, across the variety of regions and species sampled, trees that died during water shortages were less resilient to previous non-lethal droughts, relative to coexisting surviving trees of the same species. In angiosperms, drought-related mortality risk is associated with lower resistance (low capacity to reduce impact of the initial drought), while it is related to reduced recovery (low capacity to attain pre-drought growth rates) in gymnosperms. The different resilience strategies in these two taxonomic groups open new avenues to improve our understanding and prediction of drought-induced mortality.

Chilling and forcing temperatures interact to predict the onset of wood formation in Northern Hemisphere conifers.

The phenology of wood formation is a critical process to consider for predicting how trees from the temperate and boreal zones may react to climate change. Compared to leaf phenology, however, the determinism of wood phenology is still poorly known. Here, we compared for the first time three alternative ecophysiological model classes (threshold models, heat-sum models and chilling-influenced heat-sum models) and an empirical model in their ability to predict the starting date of xylem cell enlargement in spring, for four major Northern Hemisphere conifers (Larix decidua, Pinus sylvestris, Picea abies and Picea mariana). We fitted models with Bayesian inference to wood phenological data collected for 220 site-years over Europe and Canada. The chilling-influenced heat-sum model received most support for all the four studied species, predicting validation data with a 7.7-day error, which is within one day of the observed data resolution. We conclude that both chilling and forcing temperatures determine the onset of wood formation in Northern Hemisphere conifers. Importantly, the chilling-influenced heat-sum model showed virtually no spatial bias whichever the species, despite the large environmental gradients considered. This suggests that the spring onset of wood formation is far less affected by local adaptation than by environmentally driven plasticity. In a context of climate change, we therefore expect rising winter-spring temperature to exert ambivalent effects on the spring onset of wood formation, tending to hasten it through the accumulation of forcing temperature, but imposing a higher forcing temperature requirement through the lower accumulation of chilling.

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.

Water stress limits transpiration and growth of European larch up to the lower subalpine belt in an inner-alpine dry valley.

Climate change will further constrain water availability in dry inner-alpine environments and affect water relations and growth conditions in mountain forests, including the widespread larch forests. To estimate the effects of climate conditions on water balance and growth, variation in sap flow and stem radius of European larch was measured for 3 yr along an elevation transect from 1070 to 2250 m above sea level (asl) in an inner-alpine dry valley in South Tyrol/Italy. Additionally, long-term climate-growth relations were derived from tree cores. Sap flow and radial growth were reduced in dry periods up to an elevation of 1715 m, leading to maximum annual growth at 2000 m. In a wet year no growth difference between elevations was observed. Long-term tree ring data showed a positive growth response to precipitation up to 1715 m and to temperature only above 2000 m. Our results demonstrate that reduced water availability and higher atmospheric water demand limit larch at low elevation within dry Alpine regions. This indicates a general upward shift of this species' elevational amplitude upon climate change, and respective negative effects on future silvicultural use and ecosystem services at lower elevations in the European Alps.

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.

Phloem Girdling of Norway Spruce Alters Quantity and Quality of Wood Formation in Roots Particularly Under Drought.

Carbon (C) availability plays an essential role in tree growth and wood formation. We evaluated the hypothesis that a decrease in C availability (i) triggers mobilization of C reserves in the coarse roots of Picea abies to maintain growth and (ii) causes modification of wood structure notably under drought. The 6-year-old saplings were subjected to two levels of soil moisture (watered versus drought conditions) and root C status was manipulated by physically blocking phloem transport in the stem at three girdling dates (GDs). Stem girdling was done before the onset of bud break [day of the year (doy) 77], during vigorous aboveground shoot and radial stem growth (GD doy 138), and after cessation of shoot growth (GD doy 190). The effect of blockage of C transport on root growth, root phenology, and wood anatomical traits [cell lumen diameter (CLD) and cell wall thickness (CWT)] in earlywood (EW) and latewood (LW) was determined. To evaluate changes in belowground C status caused by girdling, non-structural carbohydrates (soluble sugars and starch) in coarse roots were determined at the time of girdling and after the growing season. Although fine root mass significantly decreased in response to blockage of phloem C transport, the phenology of root elongation growth was not affected. Surprisingly, radial root growth and CLD of EW tracheids in coarse roots were strikingly increased in drought-stressed trees, when girdling occurred before bud break or during aboveground stem growth. In watered trees, the growth response to girdling was less distinct, but the CWT of EW significantly increased. Starch reserves in the roots of girdled trees significantly decreased in both soil moisture treatments and at all GDs. We conclude that (i) radial growth and wood development in coarse roots of P. abies saplings are not only dependent on current photosynthates, and (ii) blockage of phloem transport induces physiological changes that outweigh drought effects imposed on root cambial activity and cell differentiation.

Robustness of xylem properties in conifers: analyses of tracheid and pit dimensions along elevational transects.

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.

Early-Warning Signals of Individual Tree Mortality Based on Annual Radial Growth.

Tree mortality is a key driver of forest dynamics and its occurrence is projected to increase in the future due to climate change. Despite recent advances in our understanding of the physiological mechanisms leading to death, we still lack robust indicators of mortality risk that could be applied at the individual tree scale. Here, we build on a previous contribution exploring the differences in growth level between trees that died and survived a given mortality event to assess whether changes in temporal autocorrelation, variance, and synchrony in time-series of annual radial growth data can be used as early warning signals of mortality risk. Taking advantage of a unique global ring-width database of 3065 dead trees and 4389 living trees growing together at 198 sites (belonging to 36 gymnosperm and angiosperm species), we analyzed temporal changes in autocorrelation, variance, and synchrony before tree death (diachronic analysis), and also compared these metrics between trees that died and trees that survived a given mortality event (synchronic analysis). Changes in autocorrelation were a poor indicator of mortality risk. However, we found a gradual increase in inter-annual growth variability and a decrease in growth synchrony in the last ∼20 years before mortality of gymnosperms, irrespective of the cause of mortality. These changes could be associated with drought-induced alterations in carbon economy and allocation patterns. In angiosperms, we did not find any consistent changes in any metric. Such lack of any signal might be explained by the relatively high capacity of angiosperms to recover after a stress-induced growth decline. Our analysis provides a robust method for estimating early-warning signals of tree mortality based on annual growth data. In addition to the frequently reported decrease in growth rates, an increase in inter-annual growth variability and a decrease in growth synchrony may be powerful predictors of gymnosperm mortality risk, but not necessarily so for angiosperms.

Cambial response of Norway spruce to modified carbon availability by phloem girdling.

We tested the hypothesis that increase in carbon (C) availability in Norway spruce saplings (Picea abies (L.) Karst.) intensifies cambial cell division and increases cell lumen diameter (CLD) and cell wall thickness (CWT) when water availability is adequate. To accomplish this, we experimentally subjected 6-year-old P. abies saplings (n = 80 trees) to two levels of soil humidity (watered versus drought conditions) and manipulated tree C status by physically blocking phloem transport at three girdling dates (GDs). Stem girdling occurred in mid-March (day of the year (doy) 77) and in mid-May (GD doy 138), i.e., ~4 weeks before the onset of bud break and during vigorous stem growth, respectively, and in early July (GD doy 190), i.e., 6 and 4 weeks after cessation of radial growth in drought-stressed trees and shoot growth in both soil humidity (SH) treatments, respectively. In response to phloem blockage a striking increase in the number of xylem cells at all GDs and reactivation of cambial activity in drought-stressed trees was detected above the girdling zone, while below girdling xylem formation stopped in both SH-treatments. Although girdling differently affected wood anatomical parameters (CLD, CWT and CLD:CWT ratio) during earlywood and latewood formation, GD had a minor effect on cambial cell division and xylem cell differentiation. Results also revealed that phloem girdling outweighed drought effects imposed on cambial activity. We explain our findings by accumulation of carbohydrates, osmotically active sugars and/or C based signaling compound(s) in response to girdling. Altogether, we conclude that wood formation in P. abies saplings is limited by C availability, which is most likely caused by high C demand belowground especially under drought.