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The metabolic activity of phytohormones and the accumulation of carbohydrates affect the reactivation of the cambial zone and the radial increment of woody plants. We aimed to monitor the dynamics of indole-3-acetic acid (IAA) concentration and amounts of soluble carbohydrates during xylem and phloem formation of one growing season (2015). Six sample trees of Scots pine (Pinus sylvestris L.), aged 80 years on average, growing in the Sobešice research site (404 m a.s.l.) in the Czech Republic were selected. We obtained microcore samples at weekly intervals by the Trephor tool method for cell formation analysis and spectrophotometric determination of IAA and soluble carbohydrate contents. We found that time of the highest concentration of IAA (last week of April) coincided with time of the maximum number of cells in the cambial zone and highest expansion of the cell enlargement stage. When the IAA concentration was too low to be measured, latewood tracheids started to form, and late phloem sieve cell formation ceased. The highest concentration of soluble carbohydrates was 200.40 ± 21.6 µg GLU per sample (May 14). This coincided with the fastest weekly xylem cell increment. This research shows that IAA and soluble carbohydrate dynamics directly affects xylem and phloem formation.

Background and aims Deciduous angiosperm trees transport xylem sap through trunks and branches in vessels within annual growth rings. Utilizing previous growth rings for sap transport could increase vessel network size and redundancy but may expose new xylem to residual air embolisms in the network. Despite the important role of vessel networks in sap transport and drought resistance, our understanding of cross-ring connections within and between species is limited. Methods We studied cross-ring connections in four temperate deciduous trees using dye staining and X-ray microcomputed tomography (microCT) to detect xylem connectivity across growth rings and quantify their impact on hydraulic conductivity. Key results Acer rubrum and Fraxinus americana had cross-ring connections visible in microCT but only A. rubrum used previous growth rings for axial sap flow. Fagus grandifolia and Quercus rubra , however, did not have cross-ring connections. Accounting for the number of growth rings that function for axial transport improved hydraulic conductivity estimates. Conclusions These data suggest that the presence of cross-ring connections may help explain aspects of whole-tree xylem sap transport and should be considered for plant hydraulics measurements in these species and others with similar anatomy. [ABSTRACT FROM AUTHOR]

Wood production is a metabolic process requiring high amount of soluble carbohydrates as source of carbon skeletons, energy, or signalling molecules. Nevertheless, the role of soluble sugars and starch has been rarely investigated in the growth to dormancy transition phase. The aim of this work was to investigate the carbohydrate concentrations and the transcription level of some candidate genes involved in cellulose and starch metabolisms in competitive carbon sinks such as phloem, maturing xylem, and xylem during the active growth to dormancy transition phase in Picea abies (L.) Karst. Stems were sampled every two weeks between July and October from potted eight-year-old Norway spruce plants. Results highlight that the cellulose biosynthesis regulated the pattern of carbohydrate changes in the phloem and maturing xylem during the transition to dormancy. In particular, the cessation of CesA (α-cellulose synthase) mRNA accumulation in maturing xylem determined a shift in the carbohydrate partitioning from the utilization sinks to the storage compartment. The content of carbohydrates was modulated in the maturing xylem by cellulose metabolism during the xylogenesis and by starch metabolism during the cambium dormancy. On the contrary, the carbon competition between utilization and storage sinks seemed to be less connected within the phloem. Since the amount of soluble carbohydrates within the maturing xylem and phloem has never been limited during the growth to dormancy transition phase, the cessation of the CesA transcription was probably determined by environmental conditions rather than by carbon limitation or sink competition. [ABSTRACT FROM AUTHOR]

The xylem of gymnosperm trees consists mainly of tracheids, which generally have larger radial diameters and thinner walls in earlywood than in latewood. Information on possible chemical differences is limited, but according to a few studies, earlywood normally contains more lignin relative to cellulose than latewood. However, for methods giving results per gram dry matter, this result is expected since a larger part of the earlywood cell wall material consists of lignin-rich middle lamella. In this study, we used Raman microspectroscopy to assess relative biopolymer composition of earlywood and latewood secondary cell walls from mature sapwood of Norway spruce (Picea abies (L.) Karst.). In addition, infrared spectroscopy of isolated earlywood and latewood from the same growth rings as well as additional growth rings was performed. Both methods showed significantly higher relative lignin content in the earlywood which confirms results from earlier studies. [ABSTRACT FROM AUTHOR]

In Larix trunks the properties of wood differ clearly radially, but the axial differences are smaller as well as being less studied. Wood anatomy is in particular poorly studied, even though all other wood properties derive from cell and tissue structure. The aim of this study was to chart variation in tracheid size (double wall thickness (2CWT), diameter of lumen (RD)) within fast grown cultivated Siberian larch (Larix sibirica Ledeb.) trunks. The differences in 2CWT and RD were clear between earlywood (EW) and latewood (LW), 2CWT increasing clearly less in EW than in LW towards the bark, while RD stayed quite stable in LW but in EW increased markedly towards the bark. The difference in 2CWT between EW and LW increased towards the upper trunk. In conclusion, the radial variation in RD and 2CWT was different between the butt and other studied heights. As the difference in 2CWT between EW and LW was smaller at the butt than the upper portion of the trunk, the wood was the most homogenous at the butt.

Ultrasonic emission (UE) testing is used to analyse the vulnerability of xylem to embolism, but the number of UEs often does not sufficiently reflect effects on hydraulic conductivity. We monitored the absolute energy of UE signals in dehydrating xylem samples hypothesizing that (i) conduit diameter is correlated with UE energy and (ii) monitoring of UE energy may enhance the utility of this technique for analysis of xylem vulnerability. Split xylem samples were prepared from trunk wood of Picea abies, and four categories of samples, derived from mature (I: earlywood, II: 30–50% latewood, III: >50% latewood) or juvenile wood (IV: earlywood) were used. Ultrasonic emissions during dehydration were registered and anatomical parameters (tracheid lumen area, number per area) were analysed from cross-sections. Attenuation of UE energy was measured on a dehydrating wood beam by repeated lead breaks. Vulnerability to drought-induced embolism was analysed on dehydrating branches by hydraulic, UE number or UE energy measurements. In split samples, the cumulative number of UEs increased linearly with the number of tracheids per cross-section, and UE energy was positively correlated with the mean lumen area. Ultrasonic emission energies of earlywood samples (I and IV), which showed normally distributed tracheid lumen areas, increased during dehydration, whereas samples with latewood (II and III) exhibited a right-skewed distribution of lumina and UE energies. Ultrasonic emission energy was hardly influenced by moisture content until ∼40% moisture loss, and decreased exponentially thereafter. Dehydrating branches showed a 50% loss of conductivity at −3.6 MPa in hydraulic measurements and at −3.9 and −3.5 MPa in UE analysis based on cumulative number or energy of signals, respectively. Ultrasonic emission energy emitted by cavitating conduits is determined by the xylem water potential and by the size of element. Energy patterns during dehydration are thus influenced by the vulnerability to cavitation, conduit size distribution as well as attenuation properties. Measurements of UE energy may be used as an alternative to the number of UEs in vulnerability analysis. [ABSTRACT FROM AUTHOR]

Four-year-old Scots pine [ Pinus sylvestris L. (Pinaceae)] seedlings were exposed to medium and severe drought stress for two consecutive years. The anatomical properties of drought-stressed Scots pine wood and their impact on the performance of destructive wood boring early instars of Hylotrupes bajulus L. (Coleoptera: Cerambycidae) were studied. Drought stress significantly decreased diameter of earlywood tracheids in both growing years and diameter of latewood tracheids after the second growing season only. Cell lumen area was significantly decreased by both medium and severe drought stress compared to well-watered controls. In addition, area of cell lumen was significantly smaller in severe drought than in medium drought treatment. The drought stress marginally increased the number of resin canals in the wood, but did not affect the size of resin canals either in wood or bark. The relative growth rate of xylophagous H. bajulus neonatal larvae was not significantly affected by drought stress during the 106-day feeding period on Scots pine wood blocks. The results show that although water availability was an important factor affecting the development and anatomy of wood cells, observed changes in wood characteristics did not affect the performance of early instars feeding on wood processed from drought-stressed young Scots pine seedlings. [ABSTRACT FROM AUTHOR]

•Seasonal wood development produces earlywood (EW) and latewood (LW) with distinct properties. The molecular mechanisms controlling EW and LW formation at different tree ages are poorly understood. •Seasonal reorganization of the xylem transcriptome was investigated in Pinus radiata at four tree ages using cDNA microarrays. Transcriptome profiles were compared with seasonal wood variation measured by SilviScan (CSIRO, Clayton, Australia). •The xylem transcriptome was considerably reorganized during seasonal change, and this reorganization showed a maturation-related pattern. The greater reorganization occurred at the transition (30%) and juvenile (21%) stages, but it declined with tree maturity (11–13%). However, this pattern does not correlate well with maturation-related patterns of seasonal wood variation. In total, 319 seasonal-responsive xylem candidate genes were identified. Many transcripts involved in primary and secondary wall biosynthesis were preferentially accumulated in EW and LW, respectively. A large proportion (45–81%) of the candidate genes are preferentially regulated at a single age and their transcript abundance may influence maturation-related patterns of seasonal wood variation. •Seasonal reorganization of the xylem transcriptome is significantly affected by tree age. Physiological changes at the transition stage may contribute to its greater seasonal transcriptome reorganization. Identified stage-preferential xylem transcripts could influence seasonal wood variation at different tree ages. [ABSTRACT FROM AUTHOR]

We studied the effects of elevated temperature and carbon dioxide concentration ([CO2]) alone and together on wood anatomy of 20-year-old Scots pine (Pinus sylvestris L.) trees. The study was conducted in 16 closed chambers, providing a factorial combination of two temperature regimes and two CO2 concentrations (ambient and elevated), with four trees in each treatment. The climate scenario included a doubling of [CO2] and a corresponding increase of 2-6 °C in temperature at the site depending on the season. Anatomical characteristics analyzed were annual earlywood, latewood and ring widths, intra-ring wood densities (earlywood, latewood and mean wood density), tracheid width, length,wall thickness, lumen diameter, wall thickness:lumen diameter ratio and mass per unit length (coarseness), and numbers of rays, resin canals and tracheids per xylem cross-sectional area. Elevated [CO2] increased ring width in four of six treatment years; earlywood width increased in the first two years and latewood width in the third year. Tracheid walls in both the earlywood and latewood tended to become thicker over the 6-year treatment period when temperature or [CO2] was elevated alone, whereas in the combined treatment they tended to become thinner relative to the tracheids of trees grown under ambient conditions. Latewood tracheid lumen diameters were larger in all the treatments relative to ambient conditions over the 6-year period, whereas lumen diameters in earlywood increased only in response to elevated [CO2] and were 3-6% smaller in the treatments with elevated temperature than in ambient conditions. Tracheid width, length and coarseness were greater in trees grown in elevated than in ambient temperature. The number of resin canals per mm² decreased in the elevated [CO2] treatment and increased in the elevated temperature treatments relative to ambient conditions. The treatments decreased the number of rays and tracheids per mm² of cross-sectional area, the greatest decrease occurring in the elevated [CO2] treatment. It seemed that xylem anatomy was affected more by elevated temperature than by elevated [CO2] and that the effects of temperature were confined to the earlywood. [ABSTRACT FROM AUTHOR]

The air-seeding hypothesis predicts that xylem embolism resistance is linked directly to bordered pit functioning. We tested this prediction in trunks, roots, and branches at different vertical and radial locations in young and old trees of Pseudosuga menziesii. Dimensions of bordered pits were measured from light and scanning electron micrographs, and physiological data were from published values. Consistent with observations, calculations showed that earlywood tracheids were more resistant to embolism than latewood tracheids, mainly from earlywood having stretchier pit membranes that can distend and cover the pit aperture. Air seeding that occurs in earlywood appears to happen through gaps between the torus edge and pit border, as shown by the similar calculated pressures required to stretch the membrane over the pit aperture and to cause embolism. Although bordered pit functioning was correlated with tracheid hydraulic diameter, pit pore size and above all pit aperture constrained conductivity the most. From roots to branches and from the trunk base to higher on the trunk, hydraulic resistance of the earlywood pit membrane increased significantly because of a decrease in the size of the pit aperture and size and number of margo pores. Moreover, overall wood conductivity decreased, in part due to lower pit conductivity and a decrease in size and frequency of pits. Structural and functional constraints leading to the trade-off of efficiency against safety of water transport were also demonstrated at the individual pit level, with a positive correlation between pit membrane resistance on an area basis and the pressure differential required to cause membrane stretching, a characteristic that is essential for pit aspiration. [ABSTRACT FROM AUTHOR]

Plants exhibit morphological and anatomical adaptations to cope the environmental constraints of their habitat. How can mechanisms for adapting to contrasting environmental conditions change the patterns of tree rings formation? In this study, we explored differences in climatic conditions of permafrost and non-permafrost zones and assessed their influence on radial growth and wood traits of Larix gmelinii Rupr (Rupr) and Larix sibirica L., respectively. We quantified the contribution of xylem cell anatomy to the tree-ring width variability. Comparison of the anatomical tree-ring parameters over the period 1963–2011 was tested based on non-parametric Mann-Whitney U test. The generalized linear modeling shows the common dependence between TRW and the cell structure characteristics in contrasting environments, which can be defined as non-specific to external conditions. Thus, the relationship between the tree-ring width and the cell production in early- and latewood are assessed as linear, whereas the dependence between the radial cell size in early- and latewood and the tree-ring width becomes significantly non-linear for both habitats. Moreover, contribution of earlywood (EW) and latewood (LW) cells to the variation of TRW (in average 56.8% and 24.4% respectively) was significantly higher than the effect of cell diameters (3.3% (EW) and 17.4% (LW)) for the environments. The results show that different larch species from sites with diverging climatic conditions converge towards similar xylem cell structures and relationships between xylem production and cell traits. The work makes a link between climate and tree-ring structure, and promotes a better understanding the anatomical adaptation of larch species to local environment conditions. [ABSTRACT FROM AUTHOR]

Research Highlights: This study emphasized the importance of multi-parameter analyses along ecological gradients for a more holistic understanding of the complex mechanism of tree-ring formation. Background and Objectives: The analysis of climatic signals from cell anatomical features measured along series of tree-rings provides mechanistic details on how environmental drivers rule tree-ring formation. However, the processes of cell development might not be independent, limiting the interpretation of the cell-based climatic signal. In this study, we investigated the variability, intercorrelations and climatic drivers of wood anatomical parameters, resulting from consequent cell developmental processes. Materials and Methods: The study was performed on thin cross-sections from wood cores sampled at ~1.3 m stem height from mature trees of Pinus sylvestris L. growing at five sampling sites along an ecological gradient from cold and wet to hot and dry within continental Southern Siberia. Tracheid number per radial file, their diameters and wall thicknesses were measured along the radial direction from microphotographs for five trees per site. These parameters were then averaged at each site for earlywood and latewood over the last 50 tree rings to build site chronologies. Their correlations among themselves and with 21-day moving climatic series were calculated. Results: Our findings showed that wood formation was not simply the result of environmentally driven independent subprocesses of cell division, enlargement and wall deposition. These processes appear to be interconnected within each zone of the ring, as well as between earlywood and latewood. However, earlywood parameters tend to have more distinctive climatic responses and lower intercorrelations. On the other hand, there are clear indications that the mechanisms of cell division and enlargement share similar climatic drivers and are more sensitive to water limitation than the process of wall deposition. Conclusions: Indications were provided that (i) earlywood formation left a legacy on latewood formation, (ii) cell division and enlargement shared more similar drivers between each other than with wall deposition, and (iii) the mechanism of cell division and enlargement along the gradient switch from water to heat limitations at different thresholds than wall deposition. [ABSTRACT FROM AUTHOR]

Research highlights: Annual wood anatomy (xylem) aids our understanding of mature wood formation and the growth strategies of trees. Background and Objectives: Catalpa bungei is an important native species in China that produces excellent quality wood. Herein, we clarified the effects of the genetic origin and cambial age on the anatomical characteristics of C. bungei wood. Materials and Methods: Six new 13-year-old C. bungei clones: '1-1' (n trees = 3), '1-3' (n trees = 3), '2-7' (n trees = 3), '2-8' (n trees = 3), '8-1' (n trees = 4), and '9-1' (n trees = 3) were removed for study from a plantation in Tianshui City, Gansu province, China. Xylem features were observed and the anatomical variables were manually measured via image analysis on (macro- micro-, and ultra-) features cut from radial increments of earlywood and latewood sampled at breast height. Results: Between the age of 1 and 2 years, wood was diffuse-porous; between the age of 3 and 9 years, wood was semi-ring-porous; and between the age of 10 and 13 years, wood was ring-porous. The effect of clones on anatomical characteristics was significant except for the microfibril angle in latewood and ring width. The transition between juvenile and mature wood was between 7 and 8 years based on patterns of radial variation in fiber length (earlywood) and microfibril angle. From the pith to the bark, fiber length, double wall thickness, fiber wall: lumen ratio, vessel diameter in earlywood, proportion of vessel in earlywood, and axial parenchyma in latewood increased significantly, whereas ring width, earlywood vessels, and the proportion of fiber decreased significantly. In addition, other features, such as vessel length, microfibril angle, and ray proportion, did not differ significantly from the pith to the bark. Conclusions: Breeding program must consider both clone and cambial age to improve the economic profitability of wood production. [ABSTRACT FROM AUTHOR]

Studying how cambial age and axial height affects wood anatomical traits may improve our understanding of xylem hydraulics, heartwood formation and axial growth. Radial strips were collected from six different heights (0–11.3 m) along the main trunk of three Manchurian catalpa (Catalpa bungei) trees, yielding 88 samples. In total, thirteen wood anatomical vessel and fiber traits were observed usinglight microscopy (LM) and scanning electron microscopy (SEM), and linear models were used to analyse the combined effect of axial height, cambial age and their interaction. Vessel diameter differed by about one order of magnitude between early- and latewood, and increased significantly with both cambial age and axial height in latewood, while it was positively affected by cambial age and independent of height in earlywood. Vertical position further had a positive effect on earlywood vessel density, and negative effects on fibre wall thickness, wall thickness to diameter ratio and length. Cambial age had positive effects on the pit membrane diameter and vessel element length, while the annual diameter growth decreased with both cambial age and axial position. In contrast, early- and latewood fiber diameter were unaffected by both cambial age and axial height. We further observed an increasing amount of tyloses from sapwood to heartwood, accompanied by an increase of warty layers and amorphous deposits on cell walls, bordered pit membranes and pit apertures. This study highlights the significant effects of cambial age and vertical position on xylem anatomical traits, and confirms earlier work that cautions to take into account xylem spatial position when interpreting wood anatomical structures, and thus, xylem hydraulic functioning. [ABSTRACT FROM AUTHOR]

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