Vessel occlusion through tyloses or gums is a natural phenomenon occurring with aging and heartwood formation, and in sapwood in response to vessel embolism. These types of vessel occlusion play a crucial role to limit the spread of pathogens and wood decay organisms, also as part of compartmentalization after wounding. In the sapwood, they can be considered to be an effective stress response.Here we review the literature on tyloses and gums in hardwoods, starting with the detailed 19th century account on tyloses by Hermine von Reichenbach. The structural diversity of tyloses (from thin-walled to sclerotic) and gum deposits is highlighted and illustrated. Our understanding of the development of vessel occlusions through vessel contact cells of the ray and axial parenchyma has greatly increased over the last decades, also thanks to ultrastructural and immunocytological studies. We critically discuss the postulated relationships between vessel-to-ray parenchyma pit size and vessel size and the incidence of either tyloses or gums and review the occurrence of these types of vessel occlusions in extant and fossil dicots. All factors identified in the literature as stimuli for vessel occlusion probably act through vessel embolism as a single direct trigger. Attempts in the literature to relate vessel occlusion with mechanisms of vessel refilling and embolism repair are controversial and invite more experimental research.

Mediterranean trees have patterns of cambial activity with one or more pauses per year, leading to intra-annual density fluctuations (IADFs) in tree rings. We analysed xylogenesis (January 2015–January 2016) in Pinus pinea L. and Arbutus unedo L., co-occurring at a site on Mt. Vesuvius (southern Italy), to identify the cambial productivity and timing of IADF formation. Dendrochronological methods and quantitative wood anatomy were applied and enabled IADF identification and classification. We showed that cambium in P. pinea was productive throughout the calendar year. From January to March 2015, post-cambial (enlarging) earlywood-like tracheids were observed, which were similar to transition tracheids. The beginning of the tree ring was therefore not marked by a sharp boundary between latewood of the previous year and the new xylem produced. True earlywood tracheids were formed in April. L-IADFs were formed in autumn, with earlywood-like cells in latewood. In A. unedo, a double pause in cell production was observed, in summer and winter, leading to L-IADFs in autumn as well. Moreover, the formation of more than one IADF was observed in A. unedo. Despite having completely different wood formation models and different life strategies, the production of earlywood, latewood and IADF cells was strongly controlled by climatic factors in the two species. Such cambial production patterns need to be taken into account in dendroecological studies to interpret climatic signals in wood from Mediterranean trees.

Mediterranean tree rings are characterized by Intra-Annual Density Fluctuations (IADFs) due to partly climate-driven cambial activity. IADFs are used as structural signals to gain information on relations between environmental conditions and eco-physiological processes during xylogenesis, with intra-annual resolution. To reach an unbiased synchronization of the IADF position within tree rings and seasonal fluctuations in environmental conditions, it is necessary to know the timing of cambial activity and wood formation, which are species- and site-specific processes. We applied the microcoring technique to analyze xylogenesis in Pinus halepensis and Arbutus unedo. To the best of our knowledge, this is the first attempt to study xylogenesis in a hardwood species forming frequent IADFs. Both species co-occur at a site in southern Italy characterized by a Mediterranean climate. To facilitate tree-ring dating and identification of IADFs, we performed traditional dendroecological analysis. We analyzed xylogenesis during summer, which is considered a constraint for xylogenesis and a trigger for IADF formation. We followed the different phases of cell development in the current wood increment with the aim of evaluating whether and which type of IADFs were formed. We additionally analyzed the same phases again in September and in winter to verify the possible formation of IADFs in fall and whether cell production and differentiation was completed by the end of the calendar year. Both species formed the same type of IADFs (earlywood-like cells within latewood), due to temporary growth restoration triggered by rain events during the period of summer drought. At the end of the calendar year, no cells in the phases of enlargement and secondary cell wall deposition occurred. A. unedo was more sensitive than P. halepensis because IADFs were formed earlier in the season and were more frequent in the tree-ring series. The dendro-anatomical approach, combining analysis of tree-ring series and of xylogenesis, helped to detect the period of IADF formation in the two species. Results are discussed in functional terms, highlighting the environmental conditions triggering IADFs, and also in methodological terms, evaluating the applicability of xylogenesis analysis in Mediterranean woods, especially when the formation of IADFs is not uniform around the stem.

Research Highlights: The antifungal assay confirmed that knotwood extractives of Scots pine inhibit the growth of wood decay fungi. Heartwood extracts of black locust were found to be much stronger free radical scavengers than the extracts of Scots pine. The extracts were deposited in the lumina and on the wall surface of cells in the impregnated sapwood. Impregnation of the sapwood blocks with Scots pine and black locust extracts reduced the fungal decay of wood. Objectives: Hydrophilic extracts of Scots pine knotwood and black locust heartwood were chemically analyzed, tested for antifungal and antioxidant properties and used for impregnation of beech and Scots pine sapwood. Materials and Methods: Scots pine knotwood and black locust heartwood were extracted, and obtained hydrophilic extractives were chemically analyzed. Extracts were analyzed for antifungal properties with the in vitro well-diffusion method. The free radical scavenging activity of wood extracts was measured colorimetrically. The retention of the extracts in the impregnated sapwood blocks was evaluated with microscopy and gravimetry. A decay test was performed with the mini block test. Results: Almost half of both Scots pine knotwood and black locust heartwood hydrophilic extracts obtained were described by phenolic compounds. The extracts were deposited in the lumina of cells and on the cell wall surface. Extractives of Scots pine knotwood had good inhibitory properties against white- and brown-rot fungi. On the other hand, extractives of black locust heartwood were found to be good radical scavengers, better than knotwood extractives of Scots pine. The extracts of Scots pine knotwood and black locust reduced the fungal decay of the tested sapwood blocks. Conclusions: The results of this research show that the less-valued knotwood of Scots pine and heartwood of black locust are a potential source of antifungal and antioxidant agents for bio-based wood preservatives.

Determining the effect of a changing climate on tree growth will ultimately depend on our understanding of wood formation processes and how they can be affected by environmental conditions. In this context, monitoring intra-annual radial growth with high temporal resolution through point dendrometers has often been used. Another widespread approach is the microcoring method to follow xylem and phloem formation at the cellular level. Although both register the same biological process (secondary growth), given the limitations of each method, each delivers specific insights that can be combined to obtain a better picture of the process as a whole. To explore the potential of visualizing combined dendrometer and histological monitoring data and scrutinize intra-annual growth data on both dimensions (dendrometer → continuous; microcoring → discrete), we developed DevX (Dendrometer vs. Xylogenesis), a visualization application using the “Shiny” package in the R programming language. The interactive visualization allows the display of dendrometer curves and the overlay of commonly used growth model fits (Gompertz and Weibull) as well as the calculation of wood phenology estimates based on these fits (growth onset, growth cessation, and duration). Furthermore, the growth curves have interactive points to show the corresponding histological section, where the amount and development stage of the tissues at that particular time point can be observed. This allows to see the agreement of dendrometer derived phenology and the development status at the cellular level, and by this help disentangle shrinkage and swelling due to water uptake from actual radial growth. We present a case study with monitoring data for Acer pseudoplatanus L., Fagus sylvatica L., and Quercus robur L. trees growing in a mixed stand in northeastern Germany. The presented application is an example of the innovative and easy to access use of programming languages as basis for data visualization, and can be further used as a learning tool in the topic of wood formation and its ecology. Combining continuous dendrometer data with the discrete information from histological-sections provides a tool to identify active periods of wood formation from dendrometer series (calibrate) and explore monitoring datasets.

Microwave modification can increase the permeability of wood by delaminating and rupturing its anatomical microstructures at their weak points. A high degree of intensity of microwave modification can cause significant structural damage to the microstructures of wood, resulting in poorer strength properties. The objective of this study was to evaluate the changes in the anatomical structure of Norway spruce (Picea abies (L.) Karst.) heartwood and sapwood after microwave modification in order to develop the most effective treatment in terms of applied energy without causing significant structural damage. Analysis with light and scanning electron microscopy were performed to evaluate the effect of microwave treatment for two different energy intensities, moderate and high intensity. The results indicated structural changes in the tracheid cells. Microscopy showed varying degrees of modification within the wood microstructure, with the heartwood samples showing a greater anatomical distortion compared to their sapwood counterparts. Furthermore, the samples were subjected to pycnometric density measurements, which indicated a reduction in skeletal and absolute density after microwave modification, for both high and moderate intensity treatment on sapwood and heartwood samples. With increasing microwave energy, a gradual increase in specific pore volume and porosity percentage of the samples were also detected.