Search Results (21)

Understanding whether long-term vascular cambium vitality in ancient trees reflects progressive decline or adaptive reprogramming is central to grasping woody plant longevity. We performed an integrative analysis of functional traits, transcriptome profiles, and metabolomic landscapes of cambial zone enriched from 80-, 500-, and 1000-year-old Styphnolobium japonicum trees. With increasing tree age, the vascular cambium showed fewer cell layers, reduced thickness, and lower auxin, gibberellin, and IAA/ABA ratios, whereas bark thickness, malondialdehyde, abscisic acid, jasmonic acid, and salicylic acid contents increased. Transcriptomic and metabolomic analyses revealed that differentially expressed genes and metabolites were primarily enriched in the cell cycle, phytohormone signaling, and phenylpropanoid biosynthesis pathways. Specifically, genes associated with cell division were down-regulated in millennial trees, whereas phenolic acids, flavonoids, and lignin-related metabolites significantly accumulated. Piecewise structural equation modeling suggested associations among tree age, transcription factors, structural genes, metabolites, and cambial functional traits. These results indicate that cambial senescence is not a simple linear decay but a highly coordinated remodeling process, providing crucial evidence for delayed senescence in long-lived woody species. Full article

Background/Objectives: Cytokinin and auxin are essential for vascular development in plants. This study aims to explore whether these two hormones exhibit crosstalk in the cambium, analogous to that observed in the apical meristem. Methods: Using the hybrid poplar (Populus alba × P. glandulosa clone ‘84K’), we integrated gravitropic induction with transcriptomic analysis and identified the cytokinin biosynthesis gene PagIPT5 as differentially expressed in a tension wood induction system. PagIPT5 overexpression lines were generated and assessed for growth-related phenotypes. The interaction between cytokinin and auxin was investigated via anatomical observation, cell proliferation assays, in situ PCR, and immunofluorescence detection of auxin and cytokinin. Results: Compared with the wild type, PagIPT5 overexpression lines showed growth inhibition and an auxin-deficient phenotype. High PagIPT5 expression in the vascular cambium elevated cytokinin levels while reducing auxin levels, leading to decreased cambial cell proliferation and suppressed xylem development. However, in the tension wood induction system, both auxin and cytokinin levels increased in the vascular cambium of tension wood relative to opposite wood. Treatment with a superoxide anion activator promotes the accumulation of both auxin and cytokinin in 84K plants. Conclusions: These results revealed an antagonistic interaction between the two hormones in the cambium zone. However, this antagonism is attenuated in tension wood, which may be induced by the accumulation of superoxide anion in tension wood. Full article

Climatic variation is inherently linked with tree phenology; however, phenological triggers depend on species and habitat. We analyzed key climatic factors for the onset of secondary growth for Scots pine (Pinus sylvestris L.) at the southern limit of its distribution in Siberia. From direct observations of developing tree rings, seasonal curves of the number of cells in the cambial zone, in the cell-expansion zone, and the total number of xylem tracheids were developed over seven years with a wide variety in the phenological dates. We found that later and shorter intervals of these stages of xylogenesis were compensated by higher maximums of kinetics curves, probably due to higher temperatures and daylengths during the respective phenophases. Air temperature and soil temperature at a depth of 20 cm converged to values (mean ± SE) 6.6 ± 0.9 °C (air) and 3.7 ± 0.4 °C (soil) for a 15-day interval prior to cambial activity onset. Date of Tsoil ≥ 3.5 °C was most closely related to cambial activity onset (r = 0.99) and preceded it by 9.6 ± 1.1 days. Cumulative temperature sums were less reliable. Apparently, both air and soil temperature thresholds have to be reached for cambial division to start in this species-habitat combination. Late abundant snowfall can yield divergence between air and soil temperatures and delay the onset of xylogenesis. Full article

Wildfires significantly affect wood properties and usability, yet their impact on hardwood species remains insufficiently understood. This study presents an exploratory characterization of moderately fire-affected pedunculate oak (Quercus robur L.) wood, combining physical, mechanical, chemical, and thermal analyses to evaluate depth-related changes within outer stem zones. Samples were collected from bark and from wood originating approximately 1 cm and 1–2 cm beneath the cambial region to evaluate radial variation associated with moderate surface fire exposure. The oven-dry density of fire-affected wood reached 720 kg·m⁻³, corresponding to values marginally below the literature reference ranges reported for unaffected oak wood. Bending strength decreased to 85.56 MPa, while compressive strength remained within or marginally above the literature reference (71.16 MPa), and Brinell hardness (42.75 MPa) stayed within the typical range for oak. Chemical and elemental analyses revealed degradation of polysaccharides and carbon enrichment in surface layers. FTIR and DSC analyses suggested partial hemicellulose degradation, structural modification of cellulose, and reduced thermal reactivity in outer stem regions. Despite these changes, the higher heating value (19.09–19.56 MJ·kg⁻¹) remained within the literature reference ranges reported for oak wood. The results suggest that under moderate surface fire conditions, fire-induced changes were primarily concentrated in outer stem layers, while inner wood retained properties comparable to the literature reference values for unaffected oak wood. These findings indicate that moderately fire-affected oak wood may remain suitable for selected material or energy-related applications following appropriate quality assessment and removal of thermally altered surface zones. Full article

Urban trees are increasingly exposed to persistent anthropogenic drivers that extend beyond climatic forcing and fundamentally alter the conditions of secondary growth. While climatic controls of cambial phenology and xylogenesis are well established, the mechanisms by which non-climatic drivers regulate cambial activity and wood formation remain fragmented and are often inferred only indirectly. Here, we develop a cambium-centred framework to synthesise current evidence on how anthropogenic drivers shape wood formation in urban and peri-urban trees. To our knowledge, this is among the first syntheses explicitly linking anthropogenic drivers to distinct stages of xylogenesis. Anthropogenic drivers are typically chronic, spatially heterogeneous, and temporally decoupled from seasonal climatic rhythms, and may alter cambial kinetics and generate anatomical signatures not captured by ring width alone. We evaluate major driver domains, including root-zone constraints, altered hydrology, urban microclimate, pollution, salinity, and mechanical disturbance, while also considering emerging drivers such as artificial light at night and microplastics. Evidence is stratified into three levels: direct observations, indirect physiological evidence, and mechanistic plausibility. Across driver classes, three recurrent anatomical patterns emerge: reduced conduit size under hydraulic or osmotic stress; anomalies in wall deposition under carbon limitation or oxidative stress; and pronounced circumferential heterogeneity under spatially localised forcing. Integrative approaches combining xylogenesis monitoring, quantitative wood anatomy, dendrometer observations and spatially explicit sampling are essential to disentangle anthropogenic from climatic effects and improve assessment of tree resilience. Full article

Long-term observations of the seasonal growth of Scots pine (Pinus sylvestris L.) tree rings in the arid conditions of the Khakass-Minusinsk Basin (southern Siberia) revealed that in 2024, trees had formed a tree ring with a typical intra-annual density fluctuation (IADF) in the transition wood. An analysis of the timing and causes of this wood structure anomaly was conducted using a combination of three approaches: (1) analyzing images of cross-sections of the forming tree ring throughout the season; (2) comparing the timing of anomalous cells’ differentiation with daily climate data; (3) comparing seasonal growth observations with calculated characteristics of the modeled growth rate and its derivatives: soil moisture and transpiration. We found that during the most severe heat wave and drought (from 22 June to 9 July), the last normal earlywood cells were yet expanding, IADF cells were being produced in the cambial zone, and the first of them began expansion, while normal cells began being produced again immediately after the subsiding of environmental stress. Apparently, low soil moisture and very high temperatures mainly impacted cells in the cambial zone, marking it as the primary target of external factors influencing tree-ring formation and structure, which is important for dendroclimatology and digital wood anatomy. This result is supported by both indirect and limited direct evidence from other sources. Full article

Stand density is one among a multitude of factors impacting the growth of trees and their responses to climatic variables, but its effect on wood quality at the scale of anatomical structure is hardly investigated. Therefore, we analyzed the radial growth and wood structure of Siberian spruce (Picea obovata Ledeb.) and Scots pine (Pinus sylvestris L.) in an experimental conifer plantation with a wide gradient of stand density in the Siberian southern taiga. The measured and indexed chronologies of the tree-ring width (TRW), number of tracheid cells per radial row in the ring produced in the cambial zone (N), cell radial diameter (D), and cell wall thickness (CWT) demonstrated the influence of the planting density. The TRW and N have a negative allometric dependence on the stand density (R² = 0.75–0.88), likely due to competition for resources. The consistent negative dependence of the D on the stand density (R² = 0.85–0.97) is log-linear and also seems to be related to tree size, while the CWT is not significantly dependent on the stand density. These findings can be used as insights in regulating cellular structure and procuring desired wood quality by silvicultural means. Both conifer species have similar climatic reactions. We observed significant suppression of TRW and D related to water deficit in May–July (both species), as well as frosty (more for pine) and low-snow (for spruce) conditions in winters, as shown by both dendroclimatic correlation and pointer year analysis. Temporal shifts in the climatic responses indicate later transition to latewood and growth cessation in sparse stands, especially in spruce. Better performance was observed in sparce and medium-density stands for both species. Full article

Heartwood formation is an important ontogenetic stage in Scots pine (Pinus sylvestris L.). The amount of heartwood determines the proportion of functionally active sapwood in the total trunk biomass as well as the quality of wood. The key criterion for heartwood formation is the death of xylem ray parenchyma cells. Previously, models that described the patterns of heartwood formation, depending on the cambial age, were derived from Scots pine trees of different ages. The cambial age is the number of annual xylem layers at the core sampling site at a certain trunk height. We studied the features of the occurrence of programmed cell death (PCD) processes during the xylem differentiation and heartwood formation of 80-year-old Scots pine trees, depending on the cambial age, under the lingonberry pine forest conditions in the transition area of the northern taiga subzone and tundra. We have shown that the distance from the cambial zone to the heartwood boundary does not change significantly with stem height. As the cambial age increases, the lifespan of the formed xylem ray parenchyma cells increases and the activity of PCD genes decreases during the formation of both (1) xylem (in the outer layers of sapwood) and (2) heartwood (in the inner layers of sapwood and transition zone). We hypothesized that the decisive factor in the PCD initiation during heartwood formation is the distance of the xylem ray parenchyma cells from the cambial zone. The younger cambium forms wider annual increments, and therefore the xylem ray parenchyma cells in these parts of the trunk reach the distance from the cambial zone earlier, which is necessary for PCD initiation. Full article

The vascular cambium is an extensive and permanent secondary meristem with wood cells products of periclinal divisions commonly contributed to two directions and arranged in radial files of trees. Cambium activity is the origin of timber production. Taxodium ascendens Brongn is an exotic species in China, and its apical meristem and cambial activity are still elusive, resulting in a lack of understanding about its wood formation and improvement. We thus addressed this knowledge gap by studying Cambium activity. For studying, twigs from five 30-year-old healthy trees were collected between February-2017 and March-2018. Anatomy deciphered its apical meristem with a Cryptomeria–Abies type. The procambium appeared after leaf primordium and initially presented five lobes as observed transversely from a one-year-old shoot. The procambium under the apical differentiated into protophloem first and then protoxylem toward the inside. It means that protoxylem differentiated later than protophloem did. After dormancy, the vascular cambium began to be active, starting in early April 2017, which was later than shoot differentiation. On 25 July 2017, the cambial zone had 9–10 immature xylem cell layers. Both initiation and cessation of the xylem preceded that of the phloem. Until 10 October 2017, few immature elements were found, indicating the translation of cells from activity to dormancy. On 15 November 2017, the cambium contained 3–4 cells in radial rows, which demonstrated the dormancy of the cambium until next spring. Furthermore, immature xylem elements increased as cell layers in the cambium zone and cell fission increased. The growth pattern of T. ascendens revealed that cambial activity is highly seasonal and dependent on changes in abiotic conditions. Thus, the wood formation in the species will be significantly altered in a changing climatic pattern. These enhance our understanding of tree growth science, wood formation, wood structure, wood properties variation and wood improvement in tree breeding. Full article

WUSCHEL (WUS)-related homeobox (WOX) protein family members play important roles in the maintenance and proliferation of the stem cells in the cambium, the lateral meristem that forms all the wood structural elements. Most studies have examined the function of these genes in angiosperms, and very little was known about coniferous trees. Pine is one of the most critical forest-forming conifers globally, and in this research, we studied the distribution of WOX4, WOX13, and WOXG genes expression in Pinus sylvestris L. trunk tissues. Further, we considered the role of TDIF(CLE41/44)/TDR(PXY) signaling in regulating Scots pine cambial activity. The distribution of CLE41/44-PXY-WOXs gene expression in Scots pine trunk tissues was studied: (1) depending on the stage of ontogenesis (the first group of objects); and (2) depending on the stage of cambial growth (the second group of objects). The first group of objects is lingonberry pine forests of different ages (30-, 80-, and 180-year-old stands) in the middle taiga subzone. At the time of selection, all the trees of the studied groups were at the same seasonal stage of development: the formation of late phloem and early xylem was occurring in the trunk. The second group of objects is 40-year-old pine trees that were selected growing in the forest seed orchard. We took the trunk tissue samples on 27 May 2022, 21 June 2022, and 21 July 2022. We have indicated the spatial separation expressed of PsCLE41/44 and PsPXY in pine trunk tissues. PsCLE41/44 was differentially expressed in Fraction 1, including phloem cells and cambial zone. Maximum expression of the PsPXY gene occurred in Fraction 2, including differentiating xylem cells. The maximum expression of the PsCLE41/44 gene occurred on 27 May, when the number of cells in the cambial zone was the highest, and then it decreased to almost zero. The PsPXY gene transcript level increased from May to the end of July. We found that the highest transcript level of the PsWOX4 gene was during the period of active cell proliferation in the cambial zone, and also in the trees with the cambial age 63 years, which were characterized by the largest number of cell layers in the cambial zone. In this study, we have examined the expression profiles of genes belonging to the ancient clade (PsWOXG and PsWOX13) in stem tissues in Scots pine for the first time. We found that, in contrast to PsWOX4 (high expression that was observed during the period of active formation of early tracheids), the expression of genes of the ancient clade of the WOX genes was observed during the period of decreased cambial activity in the second half of the growing season. We found that PsWOX13 expression was shifted to Fraction 1 in most cases and increased from the phloem side, while PsWOXG expression was not clearly bound to a certain fraction. Based on the data, the role of the CLE41/44-PXY-WOX signaling module in regulating P. sylvestris cambial growth is discussed. Full article

Molecular genetic markers of various PCD (programmed cell death) variants during xylo- and phloemogenesis have been identified for the first time in Scots pine under lingonberry pine forest conditions in Northwest Russia (middle taiga subzone). PCD is a genetically determined process. Gene profiles of serine and cysteine proteases (endopeptidases), endonucleases, and metacaspases families are often considered markers of the final xylogenesis stage. In the present study, we examined the gene expression profiles of the BFN (bifunctional endonuclease) family—BFN, BFN1, BFN2, BFN3, and peptidase (cysteine endopeptidase, CEP and metacaspase, MC5) in the radial row, in addition to the vascular phloem and cambium (F1), differentiating xylem (F2), sapwood (SW), and transition zone during the active cambial growth period of uneven-aged pine trees (25-, 63- and 164-cambial age (c.a.) years old). We have shown that the expression patterns of the PCD-related genes did not depend on the cambial age but were largely determined by plant tissue type. In the radial row F1-F2-SW, we studied the activities of enzymes, including sucrose in metabolism (sucrose synthase, three forms of invertase); antioxidant system (AOS) enzymes (superoxide dismutase, catalase); and peroxidase andpolyphenol oxidase, which belonged to AOS enzymes and were involved in the synthesis of phenolic components of cell walls. The activity of the enzymes indicated that the trunk tissues of pine trees had varying metabolic status. Molecular genetic PCD regulation mechanisms during xylem vascular and mechanical element formation and parenchyma cells’ PCD during the formation of Scots pine heartwood were discussed. Full article

Repetitive observations (direct measurements) of seasonal kinetics of xylogenesis and simulations (proxy data) with tree growth models are the two main approaches available to assess tree-ring growth and development. Both have drawbacks: short cover period for observations; limited accuracy of simulations depending on input data for models. We proposed an implementation of both approaches on the same trees to find ways for compensation. Cell numbers at subsequent xylogenesis stages were observed for Pinus sylvestris L. over five seasons in moisture-deficient habitats of Southern Siberia. The Vaganov–Shashkin model was parameterized for species and soil-landscape conditions to fit local tree-ring width chronologies (R = 0.56–0.73). Seasonal kinetics variables were then compared among themselves and with the simulated environmentally driven growth rate. The number of cells in the cambial and cell enlargement zone closely followed the curve of the 15-day moving average of the simulated growth rate (R = 0.56–0.87 at one site and R = 0.78–0.89 after shifting rate curve forward by 17–20 days at another site). The maximum number of cambium cells, which occurred within three weeks of the summer solstice, was found to be positively related with the number of tracheids in the complete tree ring (R² = 0.12–0.75 for individual seasons and 0.49 for total dataset), making it a promising short-term forecast variable for tree radial growth and productivity. Full article

Auxin has been shown to exhibit a striking concentration gradient distribution in radial sections of angiosperm and gymnosperm species, in which peak auxin levels are concentrated in dividing cambial cells, while the absolute auxin concentration sharply declines toward developing secondary phloem and xylem regions. The coincidence of auxin concentration gradient across shoot tissues and xylem cell developmental gradient has prompted that auxin could act as “a plant morphogen” to provide a positional signal for cambial cell development. However, the specific location of vascular cambium and the lack of mutants altering auxin distribution in shoots of woody species made further verification experiments difficult to explore. To address this issue, different concentrations of exogenous IAA were applied to decapitated Chinese fir (Cunninghamia lanceolata (Lamb.) Hook.) trunks in this study to induce the change in the auxin concentration gradient in radial stems, and its effects on cambial activities were examined on the physiological, cellular and molecular levels. Our findings manifested that exogenous IAA treatments resulted in vast changes in endogenous hormone concentrations (including IAA, ZR, GA₃ and ABA), cambial cell developmental behaviors and transcriptional activities of genes related to polar auxin transport (PAT), auxin signaling, the biosynthesis and signal transduction of other plant hormones and the genetic control of cambial activity. Based on above findings, we postulated a model of auxin concentration gradient involved in the control of cambial activity and secondary growth in tree trunks. In this model, the contrasting expression of AUX1/LAX and PIN family carriers in distinct Chinese fir wood-forming tissues dynamically modulates PAT into the cambial zone adjacent to the secondary phloem side and secondary xylem tissues, resulting in a sharp and wide auxin spatial gradient distribution across shoots in different stages of secondary growth, respectively. This change in auxin concentration gradient distribution in radial sections in turn acts on cambial developmental behaviors by modulating the expression of auxin signaling genes and key transcription factors and the production of other plant hormones in distinct woody tissues. Findings in this study provide important insights for understanding the biological significance of auxin concentration gradient existing in the radial stems of woody species. Full article

The balance between cell proliferation and differentiation into other cell types is crucial for meristem indeterminacy, and both growth aspects are under genetic control. The peptide-receptor signaling module regulates the activity of the cambial stem cells and the differentiation of their derivatives, along with cytokinins and auxin. We identified the genes encoding the signaling module CLE41-PXY and the regulator of vascular cambium division WOX4 and studied their expression during the period of cambial growth in the radial row: the conducting phloem/cambial zone and the differentiating xylem in two forms of Betula pendula, silver birch and Karelian birch. We have shown that the expression maximum of the BpCLE41/44a gene precedes the expression maximum of the BpPXY gene. Non-figured Karelian birch plants with straight-grained wood are characterized by a more intensive growth and the high expression of CLE41/44-PXY-WOX4. Figured Karelian birch plants, where the disturbed ratio and spatial orientation of structural elements characterizes the wood, have high levels of BpWOX4 expression and a decrease in xylem growth as well as the formation of xylem with a lower vessel density. The mutual influences of CLE41-PXY signaling and auxin signaling on WOX4 gene activity and the proliferation of cambium stem cells are discussed. Full article

The processes of cambial activity and secondary xylem and phloem differentiation are completely dependent on the influx of photoassimates. Trunk girdling is a frequently used method for studying cambial growth under conditions of different assimilate supply. We girdled 20-year-old birch trees (Betula pendula Roth) and took samples 1 cm (AG1) and 35 cm above the girdle (AG35). Tissues of ungirdled trees served as a control. A sharp increase in carbohydrates level (AG1) inhibited xylogenesis and stimulated phloemogenesis. A moderate increase (AG35) also stimulated phloemogenesis; however, xylogenesis continued. The activity of the APL gene encoding a phloem-specific transcription factor correlates with the active phloemogenesis, as it was 2.18 (AG1) and 3 (AG35) times higher than in the control. The SUC gene encoding the transmembrane sucrose transporter was up-regulated in the AG1 and AG35 zones by 2.24 and 2.51 times, respectively, compared with the control, which indicates an active sucrose loads into the cells and correlates with the preferential differentiation of parenchyma. The activity of the PIN1 gene encoding the auxin transporter was highest in zone AG35 (2.1 times higher than in the control). In zone AG1, the PIN1 activity was 1.7 times lower than that of AG35, which corresponds to the impaired differentiation of the phloem sieve tubes. The data obtained can be useful for a better understanding of physiological processes and predicting changes in the forest’s productivity under conditions of a changing climate. Full article