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.

Post-fire effects on development of leaves and secondary vascular tissues in Quercus pubescens.

An increased frequency of fire events on the Slovenian Karst is in line with future climate change scenarios for drought-prone environments worldwide. It is therefore of the utmost importance to better understand tree-fire-climate interactions for predicting the impact of changing environment on tree functioning. To this purpose, we studied the post-fire effects on leaf development, leaf carbon isotope composition (δ13C), radial growth patterns and the xylem and phloem anatomy in undamaged (H-trees) and fire-damaged trees (F-trees) of Quercus pubescens Willd. with good resprouting ability in spring 2017, the growing season after a rangeland fire in August 2016. We found that the fully developed canopy of F-trees reached only half of the leaf area index values measured in H-trees. Throughout the season, F-trees were characterized by higher water potential and stomatal conductivity and achieved higher photosynthetic rates compared to unburnt H-trees. The foliage of F-trees had more negative δ13C values than those of H-trees. This reflects that F-trees less frequently meet stomatal limitations due to reduced transpirational area and more favourable leaf-to-root ratio. In addition, the growth of leaves in F-trees relied more on the recent photosynthates than on reserves due to the fire disturbed starch accumulation in the previous season. Cambial production stopped 3 weeks later in F-trees, resulting in 60 and 22% wider xylem and phloem increments, respectively. A novel approach by including phloem anatomy in the analyses revealed that fire caused changes in conduit dimensions in the early phloem but not in the earlywood. However, premature formation of the tyloses in the earlywood vessels of the youngest two xylem increments in F-trees implies that xylem hydraulic integrity was also affected by heat. Analyses of secondary tissues showed that although xylem and phloem tissues are interlinked changes in their transport systems due to heat damage are not necessarily coordinated.

Effect of soil water availability on intra-annual xylem and phloem formation and non-structural carbohydrate pools in stem of Quercus pubescens.

Non-structural carbohydrates (NSCs, i.e., starch and soluble sugars) are frequently quantified in the context of tree response to stressful events (e.g., drought), because they serve as a carbon reservoir for growth and respiration, as well as providing a critical osmotic function to maintain turgor and vascular transport under different environmental conditions. We investigated the impact of soil water availability on intra-annual leaf phenology, radial growth dynamics and variation in NSC amounts in the stem of pubescent oak (Quercus pubescens Willd.). from a sub-Mediterranean region. For this purpose, trees growing at two nearby plots differing in bedrock and, consequently, soil characteristics (F-eutric cambisol on eocene flysch bedrock and L-rendzic leptosol on paleogenic limestone bedrock) were sampled. Non-structural carbohydrates were analysed in outer xylem and living phloem (separately for non-collapsed and collapsed parts). Results showed that xylem and phloem increments were 41.6% and 21.2%, respectively, wider in trees from F plot due to a higher rate of cell production. In contrast, the amount of NSCs and of soluble sugars significantly differed among the tissue parts and sampling dates but not between the two plots. Starch amounts were the highest in xylem, which could be explained by the abundance of xylem parenchyma cells. Two clear seasonal peaks of the starch amount were detected in all tissues, the first in September-November, in the period of leaf colouring and falling, and the second in March-April, i.e., at the onset of cambial cell production followed by bud development. The amounts of free sugars were highest in inner phloem + cambium, at the sites of active growth. Soil water availability substantially influenced secondary growth in the stem of Q. pubescens, whereas NSC amounts seemed to be less affected. The results show how the intricate relationships between soil properties, such as water availability, and tree performance should be considered when studying the impact of stressful events on the growth and functioning of trees.

Variations in Environmental Signals in Tree-Ring Indices in Trees with Different Growth Potential.

We analysed two groups of Quercus robur trees, growing at nearby plots with different micro-location condition (W-wet and D-dry) in the floodplain Krakovo forest, Slovenia. In the study we compared the growth response of two different tree groups to environmental variables, the potential signal stored in earlywood (EW) structure and the potential difference of the information stored in carbon isotope discrimination of EW and latewood (LW). For that purpose EW and LW widths and carbon isotope discrimination for the period 1970-2008 AD were measured. EW and LW widths were measured on stained microscopic slides and chronologies were standardised using the ARSTAN program. α-cellulose was extracted from pooled EW and LW samples and homogenized samples were further analysed using an elemental analyser and IRMS. We discovered that W oaks grew significantly better over the whole analysed period. The difference between D and W oaks was significant in all analysed variables with the exception of stable carbon isotope discrimination in latewood. In W oaks, latewood widths correlated with summer (June to August) climatic variables, while carbon isotope discrimination was more connected to River Krka flow during the summer. EW discrimination correlated with summer and autumn River Krka flow of the previous year, while latewood discrimination correlated with flow during the current year. In the case of D oaks, the environmental signal appears to be vague, probably due to less favourable growth conditions resulting in markedly reduced increments. Our study revealed important differences in responses to environmental factors between the two oak groups of different physiological conditions that are preconditioned by environmental stress. Environmental information stored in tree-ring features may vary, even within the same forest stand, and largely depends on the micro-environment. Our analysis confirmed our assumptions that separate EW and LW analysis of widths and carbon isotope discrimination provides complementary information in Q. robur dendroecology.