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Non-structural carbohydrates (NSCs) represent the primary carbon (C) reserves and play a crucial role for plant functioning and resilience. Indeed, these compounds are involved in the regulation between C supply and demand, and in the maintenance of hydraulic efficiency. NSCs are stored in parenchyma of woody organs, which is recognized as a proxy for reserve storage capacity of tree. Notwithstanding the importance of NSCs for tree physiology, their long-term regulation and trade-offs against growth were not deeply investigated. This work evaluated the long-term dynamics of mature tree reserves in stem and root, proxied by parenchyma features, and focusing on the trade off and interplay between the resources allocation in radial growth and reserves in stem and coarse root. In a Mediterranean beech forest, NSCs content, stem and root wood anatomy analysis, and eddy covariance data, were combined. The parenchyma fraction (RAP) of beech root and stem was different, due to differences in axial parenchyma (AP) and narrow ray parenchyma (nRP) fractions. However, these parenchyma components and radial growth showed synchronous inter-annual dynamics between the two organs. In beech stem, positive correlations were found among soluble sugars content and nRP, and among starch content and the AP. Positive correlations were found among Net Ecosystem Exchange (NEE) and AP of both organs. In contrast, NEE was negatively correlated to radial growth of root and stem. Our results suggest a different contribution of stem and roots to reserves storage, and a putative partitioning in the functional roles of parenchyma components. Moreover, a long-term trade-off of C allocation between growth and reserve pool was evidenced. Indeed, in case of C source reduction, trees preferentially allocate C towards reserves pool. Conversely, in high productive years, growth represents the major C sink., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Research on wood phenology has mainly focused on reactivation of the cambium in spring. In this study we investigated if summer drought advances cessation of wood formation and if it has any influence on wood structure in late successional forest trees of the temperate zone. The end of xylogenesis was monitored between August and November in stands of European beech and pedunculate oak in Belgium for two consecutive years, 2017 and 2018, with the latter year having experienced an exceptional summer drought. Wood formation in oak was affected by the drought, with oak trees ceasing cambial activity and wood maturation about 3 weeks earlier in 2018 compared with 2017. Beech ceased wood formation before oak, but its wood phenology did not differ between years. Furthermore, between the 2 years, no significant difference was found in ring width, percentage of mature fibers in the late season, vessel size and density. In 2018, beech did show thinner fiber walls, whereas oak showed thicker walls. In this paper, we showed that summer drought can have an important impact on late season wood phenology xylem development. This will help to better understand forest ecosystems and improve forest models., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

We explored the timing of spring xylogenesis and its potential drivers in homogeneous mature forest stands in a temperate European region. Three species with contrasting leaf development dynamics and wood anatomy were studied: European beech, silver birch and pedunculate oak. Detailed phenological observations of xylogenesis and leaf phenology were performed from summer 2017 until spring 2018. Cambium reactivation (CR) occurred before the buds of oak and birch were swollen, whereas these two phenological phases were concurrent for beech. On the other hand, initial earlywood vessels were fully differentiated (FDIEV) after leaf unfolding for all three species. Timing of CR was correlated to average ring-width of the last 10 years (2008-17), tree diameter and, partially, with tree age. In addition, the timing of FDIEV was correlated to tree age and previous year's autumn phenology, i.e., timing of wood growth cessation and onset of leaf senescence. Multivariate models could explain up to 68% of the variability of CR and 55% of the variability of FDIEV. In addition to the 'species' factor, the variability could be explained by ca 30% by tree characteristics and previous year's autumn phenology for both CR and FDIEV. These findings are important to better identify which factors (other than environment) can be driving the onset of the growing season, and highlight the influence of tree growth characteristics and previous year's phenology on spring wood phenology, wood formation and, potentially, forest production., (© The Author(s) 2020. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Cessation of xylem formation or wood growth (CWG) and onset of foliar senescence (OFS) are key autumn phenological events in temperate deciduous trees. Their timing is fundamental for the development and survival of trees, ecosystem nutrient cycling and the seasonal exchange of matter and energy between the biosphere and atmosphere, and affects the impact and feedback of forests to global change. A large-scale experimental effort and improved observational methods have allowed us to compare the timing of CWG and OFS for different deciduous tree species in Western Europe, particularly in silver birch, a pioneer species, and European beech, a late-succession species, at stands of different latitudes, of different levels of site fertility, for 2 years with contrasting meteorological and drought conditions, i.e., the low moderately dry 2017 and the extremely dry 2018. Specifically, we tested whether foliar senescence started before, after or concurrently with CWG. Onset of foliar senescence and CWG occurred generally between late September and early November, with larger differences across species and sites for OFS. Foliar senescence started concurrently with CWG in most cases, except for the drier 2018 and, for beech, at the coldest site, where OFS occurred significantly later than CWG. The behavior of beech in Spain, the southern edge of its European distribution, was unclear, with no CWG, but very low wood growth at the time of OFS. Our study suggests that OFS is generally triggered by the same drivers of CWG or when wood growth decreases in late summer, indicating an overarching mechanism of sink limitation as a possible regulator of the timing of foliar senescence., (© The Author(s) 2020. Published by Oxford University Press.)

Despite increased interest in the timing and dynamics of phloem formation, seasonal changes in the structure of phloem sieve elements remain largely unexplored. To understand better the dynamics of phloem formation and the functioning of sieve tubes in the youngest phloem in Fagus sylvatica L., we investigated repeatedly taken phloem samples during the growing season of 2017 by means of light microscopy, and transmission and scanning electron microscopy. Phloem formation started with the expansion of the overwintered early phloem sieve tubes adjacent to the cambium and concurrent cambial cell production. The highest phloem growth rate was observed in general 1 week after the onset of cambial cell production, whereas the transition from early to late phloem occurred at the end of May. Cambial cell production ceased at the end of July. The final width of the phloem increment was 184 ± 10 μm, with an early phloem proportion of 59%. Collapse of older phloem tissue is a progressive process, which continuously occurred during the sampling period. Collapse of early phloem sieve tubes started shortly after the cessation of cambial cell production. Prior to the onset of radial growth, late phloem from the previous year represented 80% of the total non-collapsed part; during the growth period, this percentage decreased to 20%. Differences were observed in both sieve tube ultrastructure and sieve plate geometry between the youngest and older phloem. However, sieve plates were never completely occluded by callose, suggesting that processes affecting the functionality of sieve tubes may differ in the case of regular collapse or injury. The youngest parts of the phloem increment from the previous year (i.e., previous late phloem) continue functioning for some time in the current growing season, but the two-step development of overwintered phloem cells also ensures a sufficient translocation pathway for photosynthates to the actively growing tissues., (© The Author(s) 2018. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

We investigated the dynamics of xylem differentiation processes and vessel characteristics in Fagus sylvatica L. to evaluate the plasticity of xylem structures under different environmental conditions. In 2008-10, analyses were performed on microcores collected weekly from two temperate sites: Menina planina (1200 m above sea level (a.s.l.)) and Panska reka (400 m a.s.l.). The duration between the onset and end of major cell differentiation steps and vessel characteristics (i.e., density, VD; mean diameter, MVD; mean area, MVA; and theoretic conductivity area, TCA) were analysed in the first and last quarters of the xylem rings, also in respect of local weather conditions (precipitation, temperature). Although the onset, duration and end of xylem formation phases differed between the two sites, the time spans between the successive wood formation phases were similar. Significant differences in MVD, MVA and TCA values were found between the first and last quarters of xylem increment, regardless of the site and year. Vessel density, on the other hand, depended on xylem-ring width and differed significantly between the sites, being about 30% higher at the high elevation site, in beech trees with 54% narrower xylem rings. Vessel density in the first quarter of the xylem ring showed a positive correlation with the onset of cell expansion, whereas a negative correlation of VD with the cessation of cell production was found in the last quarter of xylem increment. This may be explained by year-to-year differences in the timing of cambial reactivation and leaf development, which effect hormonal regulation of radial growth. No significant linkage between intra-annual weather conditions and conduit characteristics was found. It can thus be presumed that precipitation is not a limiting factor for xylem growth and cell differentiation in beech at the two temperate study sites and sites across Europe with similar weather conditions., (© The Author(s) 2018. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Despite its major economic importance and the vulnerability of Picea abies (L.) H. Karst. to climate change, how its radial growth at intra-annual resolution is influenced by weather conditions in forest stands with a high production capacity has scarcely been explored. Between 2009 and 2011, phenological variation in seasonal cambial cell production (CP) was analysed in adult P. abies trees from three contrasting sites, differing in altitude and latitude. The results indicate that the timing of cambial CP is a highly synchronic process within populations since in all cases the cambium simultaneously started and stopped producing xylem and phloem cells. Our results also demonstrate that the phenology of cambial CP is highly variable and plastic between years, depending on seasonal temperature and precipitation variation. Differences among sites, however, are only partially explained by different environmental (elevation and altitude) and climatic conditions, suggesting that local adaptation may also play a decisive role in the strategy of P. abies for adapting wood and phloem increments to function optimally under local conditions., (© The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

An age effect on growth trends and climate/growth relationships of trees can possibly be discovered by analysing the seasonal dynamics of xylem development. The aims of this study, therefore, were to compare xylem formation of young (43 ± 4 years) and old (162 ± 26 years) Smith fir (Abies georgei var. smithii (Viguie & Gaussen) W. C. Cheng & L. K. Fu) trees in the Sygera Mountains, south-eastern Tibetan Plateau and, to identify the association between wood formation and climate. The seasonal radial growth dynamics of young and old trees was monitored on microcores collected at weekly intervals during two growing seasons. Transverse sections through phloem, cambium and outermost xylem of 9-12 μ m thickness were observed with a light microscope under bright field and polarized light to follow the cambial activity and differentiation of the developing xylem. Young trees were characterized by an earlier onset of xylogenesis, a longer growing season and a higher growth rate, resulting in a higher number of xylem cells. Both young and old trees responded fast to changes of the minimum air temperature, confirming that this factor was dominant by controlling Smith fir growth on the south-eastern Tibetan Plateau.