Temporal and spatial variability of phloem structure in Picea abies and Fagus sylvatica and its link to climate.

Using a unique 8-year data set (2010-2017) of phloem data, we studied the effect of temperature and precipitation on the phloem anatomy (conduit area, widths of ring, early and late phloem) and xylem-ring width in two coexisting temperate tree species, Picea abies and Fagus sylvatica, from three contrasting European temperate forest sites. Histometric analyses were performed on microcores taken from tree stems in autumn. We found high interannual variability and sensitivity of phloem anatomy and xylem-ring widths to precipitation and temperature; however, the responses were species- and site-specific. The contrasting response of xylem and phloem-ring widths of the same tree species to weather conditions was found at the two Slovenian sites generally well supplied with precipitation, while at the driest Czech site, the influence of weather factors on xylem and phloem ring widths was synchronised. Since widths of mean annual xylem and phloem increments were narrowest at the Czech site, this site is suggested to be most restrictive for the radial growth of both species. By influencing the seasonal patterns of xylem and phloem development, water availability appears to be the most important determinant of tissue- and species-specific responses to local weather conditions.

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.

Climatic regulation of leaf and cambial phenology in Quercus pubescens: Their interlinkage and impact on xylem and phloem conduits.

Increased frequency and severity of stressful events affects the growth patterns and functioning of trees which adjust their phenology to given conditions. Here, we analysed environmental effects (temperature, precipitation, VPD and SWC) on the timing of leaf phenology, seasonal stem radial growth patterns, and xylem and phloem anatomy of Quercus pubescens in the sub-Mediterranean in the period 2014-2019, when various adverse weather events occurred, i.e. spring drought in 2015, summer fire in 2016 and summer drought in 2017. Results showed that the timings of leaf and cambium phenology do not occur simultaneously in Q. pubescens, reflecting different environmental and internal constraints. Although year-to-year variability in the timings of leaf and cambial phenology exists, their chronological sequence is fairly fixed. Different effects of weather conditions on different stages of leaf development in spring were observed. Common climatic drivers (i.e., negative effect of hot and dry summers and a positive effect of increasing moisture availability in winter and summer) were found to affect the widths of xylem and phloem increments with more pronounced effect on late formed parts. A legacy effect of the timing of leaf and cambial phenology of the previous growing season on the timing of phenology of the following spring was confirmed. Rarely available phloem data permitted a comprehensive insight into the interlinkage of the timing of cambium and leaf phenology and adjustment strategies of vascular tissues in Mediterranean pubescent oak to various environmental constraints, including frequent extreme events (drought, fire). Our results suggest that predicted changes in autumn/winter and spring climatic conditions for this area could affect the timings of leaf and stem cambial phenology of Q. pubescens in the coming years, which would affect stem xylem and phloem structure and hydraulic properties, and ultimately its performance.

Different Wood Anatomical and Growth Responses in European Beech (Fagus sylvatica L.) at Three Forest Sites in Slovenia.

European beech (Fagus sylvatica L.) adapts to local growing conditions to enhance its performance. In response to variations in climatic conditions, beech trees adjust leaf phenology, cambial phenology, and wood formation patterns, which result in different tree-ring widths (TRWs) and wood anatomy. Chronologies of tree ring width and vessel features [i.e., mean vessel area (MVA), vessel density (VD), and relative conductive area (RCTA)] were produced for the 1960-2016 period for three sites that differ in climatic regimes and spring leaf phenology (two early- and one late-flushing populations). These data were used to investigate long-term relationships between climatic conditions and anatomical features of four quarters of tree-rings at annual and intra-annual scales. In addition, we investigated how TRW and vessel features adjust in response to extreme weather events (i.e., summer drought). We found significant differences in TRW, VD, and RCTA among the selected sites. Precipitation and maximum temperature before and during the growing season were the most important climatic factors affecting TRW and vessel characteristics. We confirmed differences in climate-growth relationships between the selected sites, late flushing beech population at Idrija showing the least pronounced response to climate. MVA was the only vessel trait that showed no relationship with TRW or other vessel features. The relationship between MVA and climatic factors evaluated at intra-annual scale indicated that vessel area in the first quarter of tree-ring were mainly influenced by climatic conditions in the previous growing season, while vessel area in the second to fourth quarters of tree ring width was mainly influenced by maximum temperature and precipitation in the current growing season. When comparing wet and dry years, beech from all sites showed a similar response, with reduced TRW and changes in intra-annual variation in vessel area. Our findings suggest that changes in temperature and precipitation regimes as predicted by most climate change scenarios will affect tree-ring increments and wood structure in beech, yet the response between sites or populations may differ.

Growth-limiting factors and climate response variability in Norway spruce (Picea abies L.) along an elevation and precipitation gradients in Slovenia.

Norway spruce (Picea abies L.) is among the most sensitive coniferous species to ongoing climate change. However, previous studies on its growth response to increasing temperatures have yielded contrasting results (from stimulation to suppression), suggesting highly site-specific responses. Here, we present the first study that applies two independent approaches, i.e. the nonlinear, process-based Vaganov-Shashkin (VS) model and linear daily response functions. Data were collected at twelve sites in Slovenia differing in climate regimes and ranging elevation between 170 and 1300 m a.s.l. VS model results revealed that drier Norway spruce sites at lower elevations are mostly moisture limited, while moist high-elevation sites are generally more temperature limited. Daily response functions match well the pattern of growth-limiting factors from the VS model and further explain the effect of climate on radial growth: prevailing growth-limiting factors correspond to the climate variable with higher correlations. Radial growth correlates negatively with rising summer temperature and positively with higher spring precipitation. The opposite response was observed for the wettest site at the highest elevation, which positively reacts to increased summer temperature and will most likely benefit from a warming climate. For all other sites, the future radial growth of Norway spruce largely depends on the balance between spring precipitation and summer temperature.