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Cell death plays an important role in the determination of secondary xylem cell functions. Tracheary elements (TEs), such as vessel elements and tracheids, lose their organelles due to rapid autolysis after the completion of secondary wall thickening and lignification, and play an important role in water movement along the stem. In contrast, xylem axial and ray parenchyma cells (xylem parenchyma cells) remain alive for several years or longer and retain their organelles even after maturation. As a result, xylem parenchyma cells play important roles in nutrient storage, axial and radial transportation of materials, and defense responses in the stem. In addition, they are involved in the formation of heartwood, which contributes to increases in the resistance of the tree trunk to decay, as they synthesize heartwood components such as polyphenols prior to their death. The present review focuses on changes in long-lived ray parenchyma cells during heartwood formation, such as morphology and contents of organelles, gene expression, and survival rate in sapwood. This review also summarizes the differences in cell death characteristics between TEs and ray parenchyma cells. The elucidation of the cell death mechanism of ray parenchyma cells is expected to provide useful information for controlling the properties of heartwood. [ABSTRACT FROM AUTHOR]

Abstract Cell death plays an important role in the determination of secondary xylem cell functions. Tracheary elements (TEs), such as vessel elements and tracheids, lose their organelles due to rapid autolysis after the completion of secondary wall thickening and lignification, and play an important role in water movement along the stem. In contrast, xylem axial and ray parenchyma cells (xylem parenchyma cells) remain alive for several years or longer and retain their organelles even after maturation. As a result, xylem parenchyma cells play important roles in nutrient storage, axial and radial transportation of materials, and defense responses in the stem. In addition, they are involved in the formation of heartwood, which contributes to increases in the resistance of the tree trunk to decay, as they synthesize heartwood components such as polyphenols prior to their death. The present review focuses on changes in long-lived ray parenchyma cells during heartwood formation, such as morphology and contents of organelles, gene expression, and survival rate in sapwood. This review also summarizes the differences in cell death characteristics between TEs and ray parenchyma cells. The elucidation of the cell death mechanism of ray parenchyma cells is expected to provide useful information for controlling the properties of heartwood.

Abstract Acacia melanoxylon is well known as a valuable commercial tree species owing to its high-quality heartwood (HW) products. However, the metabolism and regulatory mechanism of heartwood during wood development remain largely unclear. In this study, both microscopic observation and content determination proved that total amount of starches decreased and phenolics and flavonoids increased gradually from sapwood (SW) to HW. We also obtained the metabolite profiles of 10 metabolites related to phenolics and flavonoids during HW formation by metabolomics. Additionally, we collected a comprehensive overview of genes associated with the biosynthesis of sugars, terpenoids, phenolics, and flavonoids using RNA-seq. A total of ninety-one genes related to HW formation were identified. The transcripts related to plant hormones, programmed cell death (PCD), and dehydration were increased in transition zone (TZ) than in SW. The results of RT-PCR showed that the relative expression level of genes and transcription factors was also high in the TZ, regardless of the horizontal or vertical direction of the trunk. Therefore, the HW formation took place in the TZ for A. melanoxylon from molecular level, and potentially connected to plant hormones, PCD, and cell dehydration. Besides, the increased expression of sugar and terpenoid biosynthesis-related genes in TZ further confirmed the close connection between terpenoid biosynthesis and carbohydrate metabolites of A. melanoxylon. Furthermore, the integrated analysis of metabolism data and RNA-seq data showed the key transcription factors (TFs) regulating flavonoids and phenolics accumulation in HW, including negative correlation TFs (WRKY, MYB) and positive correlation TFs (AP2, bZIP, CBF, PB1, and TCP). And, the genes and metabolites from phenylpropanoid and flavonoid metabolism and biosynthesis were up-regulated and largely accumulated in TZ and HW, respectively. The findings of this research provide a basis for comprehending the buildup of metabolites and the molecular regulatory processes of HW formation in A. melanoxylon.

A sapwood tree is a species in which the sapwood does not differ significantly from the heartwood and cannot be classified by shades of color. It is generally accepted that heartwood has a higher economic value than sapwood, but most of the studies related to heartwood formation have focused on heartwood trees, with less research on sapwood trees. In this paper, we take the sapwood tree Schima superba as the research object and analyze the physiological and biochemical changes in the process of heartwood formation by studying the anatomical structure of parenchyma cells, and then further explore the main categories of metabolites and compositional changes. The results showed that during heartwood formation, the parenchyma cells become inactive and the nucleus disappears, while at the same time, the storage substance starch is gradually degraded under the action of enzymes and transformed into secondary metabolites, which include terpenoids, phenols and alkaloids. The accumulation of white and colorless compounds in large quantities in the heartwood, which has some effect on the heartwood color, is an important reason why the heartwood in Schima superba shows normal formation but no difference in color from the sapwood. This study fills a gap in the mechanism of heartwood formation in sapwood trees. [ABSTRACT FROM AUTHOR]

Acacia melanoxylon is highly valued for its commercial applications, with the heartwood exhibiting a range of colors from dark to light among its various clones. The underlying mechanisms contributing to this color variation, however, have not been fully elucidated. In an effort to understand the factors that influence the development of dark heartwood, a comparative analysis was conducted on the microstructure, substance composition, differential gene expression, and metabolite profiles in the sapwood (SW), transition zone (TZ), and heartwood (HW) of two distinct clones, SR14 and SR25. A microscopic examination revealed that heartwood color variations are associated with an increased substance content within the ray parenchyma cells. A substance analysis indicated that the levels of starches, sugars, and lignin were more abundant in SP compared to HW, while the concentrations of phenols, flavonoids, and terpenoids were found to be higher in HW than in SP. Notably, the dark heartwood of the SR25 clone exhibited greater quantities of phenols and flavonoids compared to the SR14 clone, suggesting that these compounds are pivotal to the color distinction of the heartwood. An integrated analysis of transcriptome and metabolomics data uncovered a significant accumulation of sinapyl alcohol, sinapoyl aldehyde, hesperetin, 2′, 3, 4, 4′, 6′-peptahydroxychalcone 4′-O-glucoside, homoeriodictyol, and (2S)-liquiritigenin in the heartwood of SR25, which correlates with the up-regulated expression of CCRs (evm.TU.Chr3.1751, evm.TU.Chr4.654_667, evm.TU.Chr4.675, evm.TU.Chr4.699, and evm.TU.Chr4.704), COMTs (evm.TU.Chr13.3082, evm.TU.Chr13.3086, and evm.TU.Chr7.1411), CADs (evm.TU.Chr10.2175, evm.TU.Chr1.3453, and evm.TU.Chr8.1600), and HCTs (evm.TU.Chr4.1122, evm.TU.Chr4.1123, evm.TU.Chr8.1758, and evm.TU.Chr9.2960) in the TZ of A. melanoxylon. Furthermore, a marked differential expression of transcription factors (TFs), including MYBs, AP2/ERFs, bHLHs, bZIPs, C2H2s, and WRKYs, were observed to be closely linked to the phenols and flavonoids metabolites, highlighting the potential role of multiple TFs in regulating the biosynthesis of these metabolites and, consequently, influencing the color variation in the heartwood. This study facilitates molecular breeding for the accumulation of metabolites influencing the heartwood color in A. melanoxylon, and offers new insights into the molecular mechanisms underlying heartwood formation in woody plants. [ABSTRACT FROM AUTHOR]

Background and Aims Heartwood plays an important role in maintaining the structural integrity of trees. Although its formation has long been thought to be driven solely by internal ageing processes, more recent hypotheses suggest that heartwood formation acts as a regulator of the tree water balance by modulating the quantity of sapwood. Testing both hypotheses would shed light on the potential ecophysiological nature of heartwood formation, a very common process in trees. Methods We measured quantities of heartwood and sapwood, xylem conduits and the width and number of growth rings on 406 stems of Pericopsis elata with ages ranging from 2 to 237 years. A subset of 17 trees with similar ages but varying growth rate were sampled in a shaded (slower-growth) site and a sun-exposed (faster-growth) site. We used regression analysis and structural equation modelling to investigate the dynamics and drivers of heartwood formation. Key Results We found a positive effect of growth rate on the probability of heartwood occurrence, suggesting an earlier heartwood onset in faster-growing stems. After this onset age, heartwood area increased with stem diameter and age. Despite the similar heartwood production per unit stem diameter increment, shaded trees produced heartwood faster than sun-exposed trees. Tree age and hydraulics showed similar direct effects on heartwood and sapwood area of sun-exposed trees, suggesting their mutual role in driving the heartwood dynamics of sun-exposed trees. However, for shaded trees, only tree hydraulics showed a direct effect, suggesting its prominent role over age in driving the heartwood dynamics in limited growing conditions. The positive relationship between growth rate and maximum stomatal conductance supported this conclusion. Conclusions Heartwood area increases as the tree ages, but at a slower rate in trees where water demand is balanced by a sufficient water supply. Our findings suggest that heartwood formation is not only a structural process but also functional. [ABSTRACT FROM AUTHOR]

Acacia melanoxylon is well known as a valuable commercial tree species owing to its high-quality heartwood (HW) products. However, the metabolism and regulatory mechanism of heartwood during wood development remain largely unclear. In this study, both microscopic observation and content determination proved that total amount of starches decreased and phenolics and flavonoids increased gradually from sapwood (SW) to HW. We also obtained the metabolite profiles of 10 metabolites related to phenolics and flavonoids during HW formation by metabolomics. Additionally, we collected a comprehensive overview of genes associated with the biosynthesis of sugars, terpenoids, phenolics, and flavonoids using RNA-seq. A total of ninety-one genes related to HW formation were identified. The transcripts related to plant hormones, programmed cell death (PCD), and dehydration were increased in transition zone (TZ) than in SW. The results of RT-PCR showed that the relative expression level of genes and transcription factors was also high in the TZ, regardless of the horizontal or vertical direction of the trunk. Therefore, the HW formation took place in the TZ for A. melanoxylon from molecular level, and potentially connected to plant hormones, PCD, and cell dehydration. Besides, the increased expression of sugar and terpenoid biosynthesis-related genes in TZ further confirmed the close connection between terpenoid biosynthesis and carbohydrate metabolites of A. melanoxylon. Furthermore, the integrated analysis of metabolism data and RNA-seq data showed the key transcription factors (TFs) regulating flavonoids and phenolics accumulation in HW, including negative correlation TFs (WRKY, MYB) and positive correlation TFs (AP2, bZIP, CBF, PB1, and TCP). And, the genes and metabolites from phenylpropanoid and flavonoid metabolism and biosynthesis were up-regulated and largely accumulated in TZ and HW, respectively. The findings of this research provide a basis for comprehending the buildup of metabolites and the molecular regulatory processes of HW formation in A. melanoxylon. [ABSTRACT FROM AUTHOR]

Abstract Objectives Molecular cues linked to heartwood formation open new (complementary) perspectives to genetic breeding programs of Douglas-fir, a tree species largely cultivated in Europe for the natural durability and civil engineering properties of its wood. Data description RNAs from a single genotype of Douglas-fir, extracted from three distinct wood zones (outer sapwood, inner sapwood and transition zone) at four vegetative seasons to generate an extensive RNA-seq dataset used to apprehend the in-wood dynamic and seasonality of heartwood formation in this hardwood model species. Previously published data collected on somatic embryos of the same genotype could be merged with the present dataset to upgrade grade the Douglas-fir reference transcriptome.

Abstract Background The formation of a tree’s heartwood gives the wood properties such as natural decay resistance and aesthetic color, and often directly determines the value of wood products. Regulating the quantity and quality of heartwood is of great importance to the use of wood. However, the mechanism of heartwood formation has been poorly understood. Results Using Dalbergia odorifera as the study species, the number of starch grains, the morphology of the nuclei, the changes in the content of water and secondary metabolites were observed continuously in the radial direction of the xylem. The results show that from the outer toward inner sapwood, the starch grains are abundant, the length to diameter ratio of the nuclei is decreasing, and the morphology changes from elongated elliptical and then to round. In the outer transition zone, the starch grains begin to decrease abruptly and the nuclei shrink at a slower rate, with a radial width of approximately 2 mm. In the inner transition zone, the heartwood color begins to appear, the starch grains disappear and a few nuclei with reduced fluorescence are present, with a radial width of approximately 1 mm. Heartwood formation after complete disappearance of the nuclei. The moisture content of the heartwood is higher than that of the sapwood, and the inner transition zone is where the content rises. The secondary metabolites of the heartwood begin to accumulate in large quantities in the inner transition zone. Conclusion Based on the physiological changes of parenchyma cells in the xylem, the radial width of the transition zone of Dalbergia odorifera is clearly defined as approximately 3 mm. Both the water and secondary metabolite abrupt changes occur at the final stage of programmed cell death, and neither is a direct cause of programmed cell death in parenchyma cells.

Catalpa bungei ‘Jinsi’, a cultivar of C. bungei C. A. Mey., is valued for its heartwood with good overall mechanical properties, naturally durable and golden-yellow color. Little is known about heartwood formation in C. bungei ‘Jinsi’ trees. The behavior of starch, water, and nuclei was studied in the xylem tissue of C. bungei ‘Jinsi’ concerning aging in ray parenchyma cells. Blocks containing heartwood, golden zone, transition zone, and sapwood were collected from the stems of six C. bungei ‘Jinsi’ trees. The moisture content of the blocks was measured by oven drying. Changes in starch and nuclei in ray parenchyma were investigated in radial profiles from sapwood to heartwood blocks using microscopy and various staining techniques. The nuclear size and starch content gradually decreased to heartwood. While the horizontal distribution of moisture content of C. bungei ‘Jinsi’ was very varied, with the heartwood and golden zone being lower than sapwood but slightly higher than the transition zone. Starch grains were rare, but nuclei were still present in some ray parenchyma cells in the heartwood and golden zone. The nuclei showed irregular shape and elongation before disintegration. These results suggest that the most apparent change occurs in the transition zone, the critical location involved in forming C. bungei ‘Jinsi’ heartwood. Water and starch appear to be actively engaged in heartwood formation. The loss of function of ray parenchyma cells results from heartwood formation.

Objectives: Molecular cues linked to heartwood formation open new (complementary) perspectives to genetic breeding programs of Douglas-fir, a tree species largely cultivated in Europe for the natural durability and civil engineering properties of its wood. Data description: RNAs from a single genotype of Douglas-fir, extracted from three distinct wood zones (outer sapwood, inner sapwood and transition zone) at four vegetative seasons to generate an extensive RNA-seq dataset used to apprehend the in-wood dynamic and seasonality of heartwood formation in this hardwood model species. Previously published data collected on somatic embryos of the same genotype could be merged with the present dataset to upgrade grade the Douglas-fir reference transcriptome. [ABSTRACT FROM AUTHOR]

Background: The formation of a tree's heartwood gives the wood properties such as natural decay resistance and aesthetic color, and often directly determines the value of wood products. Regulating the quantity and quality of heartwood is of great importance to the use of wood. However, the mechanism of heartwood formation has been poorly understood. Results: Using Dalbergia odorifera as the study species, the number of starch grains, the morphology of the nuclei, the changes in the content of water and secondary metabolites were observed continuously in the radial direction of the xylem. The results show that from the outer toward inner sapwood, the starch grains are abundant, the length to diameter ratio of the nuclei is decreasing, and the morphology changes from elongated elliptical and then to round. In the outer transition zone, the starch grains begin to decrease abruptly and the nuclei shrink at a slower rate, with a radial width of approximately 2 mm. In the inner transition zone, the heartwood color begins to appear, the starch grains disappear and a few nuclei with reduced fluorescence are present, with a radial width of approximately 1 mm. Heartwood formation after complete disappearance of the nuclei. The moisture content of the heartwood is higher than that of the sapwood, and the inner transition zone is where the content rises. The secondary metabolites of the heartwood begin to accumulate in large quantities in the inner transition zone. Conclusion: Based on the physiological changes of parenchyma cells in the xylem, the radial width of the transition zone of Dalbergia odorifera is clearly defined as approximately 3 mm. Both the water and secondary metabolite abrupt changes occur at the final stage of programmed cell death, and neither is a direct cause of programmed cell death in parenchyma cells. [ABSTRACT FROM AUTHOR]

The development of stem heartwood and the factors that control it play an important role in tree physiology, thereby impacting demographic and ecological processes of woody species. We investigated the relationships of stem heartwood with site- and tree-level variables in Pinus canariensis plantations. A total of 30 plots were sampled in the island of Gran Canaria, Spain, over a large elevation range (995–1875 m) and on terrain with different slopes (4%–70%) and exposures. The 15 pines closest to each plot center were measured and cored to quantify growth rates and the size of heartwood, also known as "tea". We used generalized linear mixed models (GLMMs) to account for both fixed and random effects while evaluating the best predictors of heartwood presence. Stem diameter was the variable most correlated with heartwood radius, and allowing for a random slope and intercept of this relationship accounted for spatially related variability. Furthermore, the GLMM model became more effective when the relationship between stem diameter and heartwood was modeled using the presence/absence of the "tea" rather than its measured size. Other site- and tree-level variables either were not statistically significant or improved the model relatively little. Because stem heartwood affects both wood quality and the amount of carbon that trees can store, our findings have implications for forest management and carbon-conscious policies. [ABSTRACT FROM AUTHOR]

The purpose of this study was to investigate the relationship between lignan biosynthesis and programmed cell death (PCD) of ray parenchyma cells during the heartwood formation of Taiwania (Taiwania cryptomerioides Hayata). Since the PCD of ray parenchyma cells and the synthesis of lignans are the two main processes involved in the formation of heartwood, both of which need to be completed through gene regulation. Based on the results of genomics and bioinformatics analysis, that the PCD of tracheids are induced by genotoxic, and the PCD of ray parenchyma cells is induced by biological factors, such as fungi, bacteria, and viruses, which could induce oxidative stress. According to the results of time−of−flight secondary ion mass spectrometry (ToF−SIMS) analysis, lignans are produced in ray parenchyma cells, and the accumulation of savinin and its downstream lignans might be the cause of PCD in ray parenchyma cells. An in vitro experiment further confirmed that the accumulation of savinin could cause protoplasts of Taiwania's xylem to produce taiwanin A, which is the marker of heartwood formation in Taiwania. Resulting in an increase in reactive oxygen species (ROS) content, which could induce oxidative stress in ray parenchyma cells and potentially lead to PCD. Based on these findings, we conclude that accumulation of savinin could be induced PCD of ray parenchyma cells in heartwood formation in Taiwania. [ABSTRACT FROM AUTHOR]

R2R3-MYB transcription factors (TFs) form one of the most important TF families involved in regulating various physiological functions in plants. The heartwood of Dalbergia odorifera is a kind of high-grade mahogany and valuable herbal medicine with wide application. However, the role of R2R3-MYB genes in the growth and development of D. odorifera, especially their relevance to heartwood formation, has not been revealed. A total of 126 R2R3-MYBs were screened from the D. odorifera genome and named DodMYB1-126 based on their location on 10 chromosomes. The collinearity results showed that purification selection was the main driving force for the evolution of the R2R3-MYB TFs family, and whole genome/fragment replication event was the main form for expanding the R2R3-MYB family, generating a divergence of gene structure and function. Comparative phylogenetic analysis classified the R2R3-MYB TFs into 33 subfamilies. S3-7,10,12-13,21 and N4-7 were extensively involved in the metabolic process; S9,13,16-19,24-25 and N1-3,8 were associated with the growth and development of D. odorifera. Based on the differential transcriptional expression levels of R2R3-MYBs in different tissues, DodMYB32, DodMYB55, and DodMYB89 were tentatively screened for involvement in the regulatory process of heartwood. Further studies have shown that the DodMYB89, localized in the nucleus, has transcriptional activation activity and is involved in regulating the biosynthesis of the secondary metabolites of heartwood by activating the promoters of the structural genes DodI2'H and DodCOMT. This study aimed to comprehensively analyze the functions of the R2R3-MYB TFs and screen for candidate genes that might be involved in heartwood formation of D. odorifera. [ABSTRACT FROM AUTHOR]

Using 146 journals, 6 chapters, 4 reports, 2 books, and a proceeding, this research seeks to investigate the factors that influence variations in the proportion of heartwood. The data were collected through an online search in databases such as Scopus, Science Citation Index, Science Direct, Google Scholar, and ResearchGate with seven search queries, namely heartwood variability, heartwood proportion, tree age of heartwood, growth rate of heartwood, silvicultural treatment of heartwood, as well as genetic and environmental effects of heartwood. By eliminating papers that were out of topic or title, purpose and year of publication, 67 journal articles, 4 chapters, 3 reports, a book, and a proceeding were selected. The review results showed that the amount of heartwood was influenced by its position in the tree, tree variations, such as age, growth rate, and species/genera, as well as the growing conditions, including location and environmental factors. It is also affected by various silvicultural treatments, namely spacing, thinning, pruning, fertilization, and irrigation. These findings indicate that the amount of heartwood is more controlled by the diameter of the wood than the age. Furthermore, several studies reported that the growth rate of sapwood rings was significantly higher than the heartwood rings, which can reduce the quality of solid wood used for construction. Therefore, it is necessary to control the proportion of heartwood through genetics and environmental factors, especially genetics, which has more effects on the growth rate of tree diameter than environmental factors. Previous studies revealed that tree diameter has a weak or negative correlation with other wood properties, such as basic density, color, and wood stiffness, which are generally important in global markets. It is important for breeders to concentrate on using species with superior genetic variations as well, especially the highly demanded traits in further studies.

We investigated the distribution of extractive, phenolic, and polysaccharides content of young S. mahagoni trees aged 1 to 5 years. The wood powder materials in two vertical positions (bottom and top) were successively extracted using n-hexane, methanol, and hot water solutions. The results showed that there was a significant interaction between the tree age and axial position in all parameters. The methanol and total extractive content decreased significantly from 1 to 3-year-old at the top part. In contrast, the amount of hot water and total extractive increased significantly from 3 to 4-year-old trees at the bottom part. Furthermore, a significant increase was also found at the bottom heartwood for hexane, methanol, and the total extractive from 4 to 5-year-old trees. The reverse trend was observed in the polar fractions (methanol and hot water) at the top of the trees. Generally, the extractive pattern tends to decrease from bottom to top with the highest proportion of methanol extract (70% based on the total extractive weight), and it increased from sapwood to heartwood. The total phenolic content (TPC) and total soluble polysaccharides (TSP) generally increase with tree age (except for TSP from 2 to 3-year-old trees at the top part). The high content of TPC in the heartwood and TSP in the sapwood indicated the process of heartwood formation. In addition, a drastic decrease in the methanol extract and TSP at the age of 3 years at the top indicated that the TSP was transferred from top to bottom for the heartwood formation at the bottom part of 4-year-old trees.

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. [ABSTRACT FROM AUTHOR]

The purpose of this study was to investigate the relationship between lignan biosynthesis and programmed cell death (PCD) of ray parenchyma cells during the heartwood formation of Taiwania (Taiwania cryptomerioides Hayata). Since the PCD of ray parenchyma cells and the synthesis of lignans are the two main processes involved in the formation of heartwood, both of which need to be completed through gene regulation. Based on the results of genomics and bioinformatics analysis, that the PCD of tracheids are induced by genotoxic, and the PCD of ray parenchyma cells is induced by biological factors, such as fungi, bacteria, and viruses, which could induce oxidative stress. According to the results of time−of−flight secondary ion mass spectrometry (ToF−SIMS) analysis, lignans are produced in ray parenchyma cells, and the accumulation of savinin and its downstream lignans might be the cause of PCD in ray parenchyma cells. An in vitro experiment further confirmed that the accumulation of savinin could cause protoplasts of Taiwania’s xylem to produce taiwanin A, which is the marker of heartwood formation in Taiwania. Resulting in an increase in reactive oxygen species (ROS) content, which could induce oxidative stress in ray parenchyma cells and potentially lead to PCD. Based on these findings, we conclude that accumulation of savinin could be induced PCD of ray parenchyma cells in heartwood formation in Taiwania.

Acacia catechu (Sha)-bearing forests are the primary sources of cutch, a tannin extract from the heartwood of Sha trees. Sha forests in Myanmar are managed for cutch production, and tree harvesting for cutch is regulated by an official diameter limit (ODL, 30 cm DBH [diameter at breast height]). We explored sustainable Sha forest management for cutch production through stand inventory surveys and informal interviews with locals and forest managers. We compared Sha forests with six different official harvest histories and assessed seedlings and saplings as well as the size and species of harvested stumps and remaining trees. We found that the forest understory was disturbed by surface fire, and all Sha seedlings and saplings < 1.7 m in height showed post-fire marks. We observed a regeneration gap between 1.7 m and 2.7 m, which might indicate the flame height of the surface fire. The "illegal" harvest exceeded the official harvest; only 5% of the harvested stumps were found to be larger than the ODL. Local harvesting of cutch appeared to be limited by the stem diameter required for heartwood formation (15 cm DBH). Stump data revealed that the forests were utilized not only for cutch but also for other purposes, including fuel and timber. Despite fire and local harvesting, local forest utilization patterns appear to be reasonable, although they are illegal. Implementing fire control and community management of forests along with clear definition of property rights could help in sustainably managing Sha forests for cutch production. [ABSTRACT FROM AUTHOR]

Induction of heartwood formation in 6-year-old Indian sandalwood (Santalum album L.) trees by treatment with carbon dioxide, ethylene, nitrogen, and wounding was investigated. All treatments induced fragrant heartwood formation upward and downward from the drill hole. The amount of heartwood formed above and below the drill hole depended on the treatment in the order nitrogen>carbon dioxide>ethylene>wounding, whereas the radial extension proportion was, in order, nitrogen>carbon dioxide>ethylene=wounding. Based on the chemical analysis (GC-MS) and evaluation of the essential oil quality and heartwood properties, heartwood induced by carbon dioxide showed the maximum similarities to naturally formed heartwood, which included the same color, similar chemical composition, reasonable oil content, and quality essential oil, whereas ethylene, nitrogen, and wounding treatment showed fewer similarities to natural heartwood. The results suggest that carbon dioxide is a promising candidate gas elicitor for inducing heartwood formation in young S. album. [ABSTRACT FROM AUTHOR]

The investigation described herein discusses the morpho-anatomical characteristics of xylem parenchyma cells of European ash stems undergoing heartwood formation. The research material comprised of cross and radial sections of wood obtained from stems at breast height of 91-year-old ash trees, half of which had visible ash dieback symptoms. The radial section of the wood samples was stained with acetocarmine to detect nuclei and with I2KI solution to observe starch grains in parenchyma cells, both of radial and axial systems. Additionally, microscopic slides were stained with Alcianblue Safranin-O, and fluorescence microscopy was applied to detect lignified cell walls. The color of sapwood and heartwood distinctly differed - heartwood extracts were detected in approx. 47 rings. Most of the parenchyma cells had nuclei present in both wood zones. Also starch grains were detected in the majority of the tree rings of the researched samples. All of the xylem parenchyma cell walls of axial and radial systems were lignified. The research revealed that lignification, parenchyma cells death and the release of heartwood extracts are processes remote in time and space. Furthermore, parenchyma cell walls lignification did not figure as a sign of the upcoming parenchyma cells death. Based on the current research, ash dieback disease might slightly impact the development paths of parenchyma cells. Compared with scores reported for other trees, European ash parenchyma cells longevity is indeed remarkable. [ABSTRACT FROM AUTHOR]

This study aims to investigate the sugar components which play an important role in the process of heartwood formation in young Swietania mahagoni. The methanol-water (70/30, v/v) extract was analyzed by GC-MS to determine sugar constituents. The content of methanol-water extract ranged from 3.7 to 7.8% based on dried wood. After a 4-5 year duration of heartwood formation in the sapwood, monosaccharide and alditol fractions doubly increased. This was followed by almost five times the content of cyclitols. In a radial variation, the content of major monosaccharides including fructose, glucose, mannose and xylose decreased from sapwood to heartwood, while arabinose, galactose, and sucrose in the age 5 years were only found in the sapwood. Meanwhile, the main content of cyclitols and alditols (arabitol) significantly decreased from sapwood to heartwood. The main monosaccharides and disaccharides represented by sucrose played a key role in the process of heartwood formation. Further studies are required to investigate the phenolic and lipophilic components as both are involved in heartwood formation. In addition, similar studies on alditol and cyclitol regarding their role in tropical wood are necessary. [ABSTRACT FROM AUTHOR]

Abstract Taiwania (Taiwania cryptomerioides Hayata) has long been regarded as a living fossil from the Tertiary period of Mesozoic Era for its distinguished yellowish-red color with purplish-pink streaks presented in its heartwood. With this elegant appearance that matches the color “red” for good fortune in the Taiwanese culture, Taiwania is supposed to be a popular wood in Taiwan where it is a native species of. Extractives contribute to the properties of wood. It is a fascinating subject to investigate extractives biosynthesis in the process of heartwood formation. Up to date, there is no phytochemistry study of Taiwania sapwood. In this study, three new sesquiterpenoids, Taiwania A (1), Taiwania B (2), and Taiwania C (3), together with 75 known compounds in the Taiwania sapwood. The structures of extractives were determined by analysis of spectroscopic data and comparison with the literatures. This study supported secondary reaction lignans could be found in sapwood that confirmed our previous research on the Taiwania-type of heartwood formation.

We investigated the distribution of extractive, phenolic, and polysaccharides content of young S. mahagoni trees aged 1 to 5 years. The wood powder materials in two vertical positions (bottom and top) were successively extracted using n-hexane, methanol, and hot water solutions. The results showed that there was a significant interaction between the tree age and axial position in all parameters. The methanol and total extractive content decreased significantly from 1 to 3-year-old at the top part. In contrast, the amount of hot water and total extractive increased significantly from 3 to 4-year-old trees at the bottom part. Furthermore, a significant increase was also found at the bottom heartwood for hexane, methanol, and the total extractive from 4 to 5-year-old trees. The reverse trend was observed in the polar fractions (methanol and hot water) at the top of the trees. Generally, the extractive pattern tends to decrease from bottom to top with the highest proportion of methanol extract (70% based on the total extractive weight), and it increased from sapwood to heartwood. The total phenolic content (TPC) and total soluble polysaccharides (TSP) generally increase with tree age (except for TSP from 2 to 3-year-old trees at the top part). The high content of TPC in the heartwood and TSP in the sapwood indicated the process of heartwood formation. In addition, a drastic decrease in the methanol extract and TSP at the age of 3 years at the top indicated that the TSP was transferred from top to bottom for the heartwood formation at the bottom part of 4-year-old trees. [ABSTRACT FROM AUTHOR]

Erythrophleum fordii Oliv. (Caesalpiniaceae) is a rosewood species naturally distributed in southeast Asia and south China, and the heartwood is commonly used for high-quality crafts and furniture. While there are differences in heartwood development among single trees with distinct social status, the relationship between heartwood development and growth performance remains unclear. This information is essential to improve plantation management for high-yield heartwood production. Forty dominant, intermediate and suppressed trees were sampled from E. fordii plantations aged in 32–35 years in Pingxiang City, Guangxi, China. Stem analysis was carried out to determine horizontal and vertical variations of heartwood and sapwood. Number of annual rings, diameter and area of heartwood and sapwood as well as ratios of heartwood diameter and area at breast height were all significantly influenced by the social status of trees in stands (P < 0.05). In these mid-aged plantations, E. fordii stems developed heartwood once the xylem diameter reached 5–10 cm, and then heartwood diameter and area increased with increasing xylem diameter. Heartwood ring numbers, diameter and area as well as their ratios decreased with increasing height, while sapwood ring numbers and diameter were relatively constant within the section where heartwood occurred. Heartwood and sapwood diameters were equal at heights of approximately 6-m for dominant, 5-m for intermediate and 3-m for suppressed trees. Dominant trees differed considerably from intermediate and suppressed trees in heartwood volume, while heartwood volume ratios were all below 30%, and near 90% in the stem section below 8-m height regardless of social status. Relationship analysis showed that DBH was the most important factor influencing heartwood in even-aged stands. The findings provide evidence for crop tree selection, thinning regimes and reasonable management of plantations of E. fordii. [ABSTRACT FROM AUTHOR]

The heartwood formation process is little known in fast growing plantation woods. Therefore, the aims of this study were to determine how planting spacing and tree age affect the formation and proportion of heartwood and sapwood, as well as the density of eucalyptus wood. Trees from a eucalyptus clonal plantation (Eucalyptus grandis × E. urophylla hybrid) cultivated in three spacings (3 × 1, 3 × 2 and 3 × 3 m) were sampled at 14, 27, 32, 53, 64 and 76 months of age. Heartwood percentage was quantified with Dimethyl yellow indicator, while the wood density was determined by X-ray densitometry. The heartwood percentage, wood volume, heartwood density and wood density were correlated with the different growth rates. The heartwood formation process started between 32 and 53 months, regardless of spacing. The heartwood proportion doubled with increasing age in the widest spacing and increased about four times in 3 × 1 and 3 × 2 m spacing. The planting spacing influenced the growth rates of the trees, but did not affect the heartwood and sapwood percentage or density. The greatest increase in density values occurred between the first year of growth (14 months) and the beginning of heartwood formation (53 months). [ABSTRACT FROM AUTHOR]

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.

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Taiwania (Taiwania cryptomerioides Hayata) has long been regarded as a living fossil from the Tertiary period of Mesozoic Era for its distinguished yellowish-red color with purplish-pink streaks presented in its heartwood. With this elegant appearance that matches the color "red" for good fortune in the Taiwanese culture, Taiwania is supposed to be a popular wood in Taiwan where it is a native species of. Extractives contribute to the properties of wood. It is a fascinating subject to investigate extractives biosynthesis in the process of heartwood formation. Up to date, there is no phytochemistry study of Taiwania sapwood. In this study, three new sesquiterpenoids, Taiwania A (1), Taiwania B (2), and Taiwania C (3), together with 75 known compounds in the Taiwania sapwood. The structures of extractives were determined by analysis of spectroscopic data and comparison with the literatures. This study supported secondary reaction lignans could be found in sapwood that confirmed our previous research on the Taiwania-type of heartwood formation. [ABSTRACT FROM AUTHOR]

Formation of extractive-rich heartwood is a process in live trees that make them and the wood obtained from them more resistant to fungal degradation. Despite the importance of this natural mechanism, little is known about the deposition pathways and cellular level distribution of extractives. Here we follow heartwood formation in Larix gmelinii var. Japonica by use of synchrotron infrared images analyzed by the unmixing method Multivariate Curve Resolution – Alternating Least Squares (MCR-ALS). A subset of the specimens was also analyzed using atomic force microscopy infrared spectroscopy. The main spectral changes observed in the transition zone when going from sapwood to heartwood was a decrease in the intensity of a peak at approximately 1660 cm-1 and an increase in a peak at approximately 1640 cm-1. There are several possible interpretations of this observation. One possibility that is supported by the MCR-ALS unmixing is that heartwood formation in larch is a type II or Juglans-type of heartwood formation, where phenolic precursors to extractives accumulate in the sapwood rays. They are then oxidized and/or condensed in the transition zone and spread to the neighboring cells in the heartwood.

Formation of extractive-rich heartwood is a process in live trees that make them and the wood obtained from them more resistant to fungal degradation. Despite the importance of this natural mechanism, little is known about the deposition pathways and cellular level distribution of extractives. Here we follow heartwood formation in Larix gmelinii var. Japonica by use of synchrotron infrared images analyzed by the unmixing method Multivariate Curve Resolution – Alternating Least Squares (MCR-ALS). A subset of the specimens was also analyzed using atomic force microscopy infrared spectroscopy. The main spectral changes observed in the transition zone when going from sapwood to heartwood was a decrease in the intensity of a peak at approximately 1660 cm-1 and an increase in a peak at approximately 1640 cm-1. There are several possible interpretations of this observation. One possibility that is supported by the MCR-ALS unmixing is that heartwood formation in larch is a type II or Juglans -type of heartwood formation, where phenolic precursors to extractives accumulate in the sapwood rays. They are then oxidized and/or condensed in the transition zone and spread to the neighboring cells in the heartwood. [ABSTRACT FROM AUTHOR]

ABSTRACT The objective of this work was to evaluate the influence of different useful areas provided by the planting spacings (3 × 1, 3 × 2, 3 × 3, 3 × 4 m) on the production and permeability of heartwood and sapwood of Eucalyptus grandis and Eucalyptus grandis × Eucalyptus urophylla clones at 4 years old. Regardless of the clone, there was no effect of planting spacing on the heartwood/sapwood relation and wood permeability. All clones showed a heartwood decrease with increased height, regardless of planting spacing, and E. grandis (B) was the only one that had its heartwood percentage positively and significantly correlated with the dendrometric variables (DBH and commercial height). The use of the dymethil yellow compound was indispensable in defining the heartwood and sapwood regions in the samples from 50% of the stem height. The heartwood permeability was low in all evaluated clones, suggesting vessel obstruction by tyloses or other deposits, while sapwood permeability reached 405.4 cm3/cm.atm.s.

Catalpa bungei 'Jinsi', a cultivar of C. bungei C. A. Mey., is valued for its heartwood with good overall mechanical properties, naturally durable and golden-yellow color. Little is known about heartwood formation in C. bungei 'Jinsi' trees. The behavior of starch, water, and nuclei was studied in the xylem tissue of C. bungei 'Jinsi' concerning aging in ray parenchyma cells. Blocks containing heartwood, golden zone, transition zone, and sapwood were collected from the stems of six C. bungei 'Jinsi' trees. The moisture content of the blocks was measured by oven drying. Changes in starch and nuclei in ray parenchyma were investigated in radial profiles from sapwood to heartwood blocks using microscopy and various staining techniques. The nuclear size and starch content gradually decreased to heartwood. While the horizontal distribution of moisture content of C. bungei 'Jinsi' was very varied, with the heartwood and golden zone being lower than sapwood but slightly higher than the transition zone. Starch grains were rare, but nuclei were still present in some ray parenchyma cells in the heartwood and golden zone. The nuclei showed irregular shape and elongation before disintegration. These results suggest that the most apparent change occurs in the transition zone, the critical location involved in forming C. bungei 'Jinsi' heartwood. Water and starch appear to be actively engaged in heartwood formation. The loss of function of ray parenchyma cells results from heartwood formation., Competing Interests: The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (© 2024 The Authors.)

Five chemical elicitors––6-benzyladenine (BA), ethephon (ETH), methyl jasmonate (MeJA), hydrogen peroxide (H2O2) and calcium chloride (CaCl2)––were used to treat 1- and 5-year-old sandal trees (Santalum album L.) to assess their effects on growth, heartwood formation and concrete oil synthesis. The results showed that some newly formed branches in stems that were induced by BA and ETH displayed leaf senescence and developed new smaller and light-green leaves. The relative percentage of concrete oil from the heartwood of water-treated trees (0.65%) was significantly lower than that from trees treated with 4 mM H2O2 (2.85%) and 4 mM BA (2.75%) within one year. Four mM BA, H2O2 and CaCl2 induced a significantly higher level of sesquiterpenoids than heartwood treated with 2 mM of these elicitors. Four mM MeJA induced significantly less sesquiterpenoids than heartwood treated with 2 mM MeJA. Morphological, physiological, and chromatographic–spectrometric technologies were integrated to trace the potential function of these exogenously applied chemical elicitors. The results may have important applications and provide a better understanding of the molecular mechanism of heartwood formation and hardening in young sandalwood trees.

The transition from the living water-transporting sapwood to heartwood involves in many tree species impregnation with extractives. These differ in amount and composition, and enhance resistance against bacteria, insects or fungi. To understand the synthesis, transport and impregnation processes new insights into the biochemical processes are needed by in-situ methods. Here we show the extractive distribution in pine (Pinus sylvestris) microsections with a high lateral resolution sampled in a non-destructive manner using Confocal Raman Microscopy. Integrating marker bands of stilbenes and lipids enables to clearly track the rapid change from sapwood to heartwood within one tree ring. The higher impregnation of the cell corner, compound middle lamella, the S3 layer and pits reveals the optimization of decay resistance on the micron-level. Furthermore, deposits with changing chemical composition are elucidated in the rays and lumen of the tracheids. The spectral signature of these deposits shows the co-location of lipids and pinosylvins with changing ratios from the living to the dead tissue. The results demonstrate that the extractive impregnation on the micro- and nano-level is optimized by a symbiotic relationship of lipids and pinosylvins to enhance the tree's resistance and lifetime. [ABSTRACT FROM AUTHOR]

Changes in the morphology and functions of vacuoles provide useful information about the mechanism of cell death. In the present study, we monitored the morphology and contents of vacuoles during the death of ray parenchyma cells in the conifer Cryptomeria japonica. In differentiating xylem, ray parenchyma cells had large central vacuoles. In sapwood, vacuoles in ray parenchyma cells contained proteins, an indication that one of the main functions of these vacuoles might be protein storage. A dramatic decrease in the protein content of some vacuoles was detected in the intermediate wood before the initiation of vacuole rupture. Although vacuole rupture was detected from the intermediate wood to the outermost heartwood, some vacuoles were obviously enlarged in the inner intermediate wood. Condensed nuclei were first observed after the rupture of these large vacuoles in ray parenchyma cells. It seems plausible that the autolysis of the contents of ray parenchyma cells might be caused by the rupture of the enlarged vacuoles in the inner intermediate wood. [ABSTRACT FROM AUTHOR]

The heartwood of Taiwania (Taiwania cryptomerioides Hayata) has the excellent decay resistance and fascinating color qualities due to the accumulation of bioactive compounds. However, limited information was addressed to the heartwood formation of this species. In this study, we established the distribution of the living ray parenchyma cells and profiled the major bioactive compounds across and along the Taiwania stem. The ray parenchyma cells would die within 1-2 annual rings of the transition zone, which occurs around 8-13 annual rings counted from the cambium. The gradual loss of the starch grains accompanied with the deposition of the colored materials into the cell lumens started from the outer transition zone. The radial distribution of the extractive contents and its signature bioactive compounds across Taiwania stems gradually increased from the transition zone toward the heartwood tissues. The aforementioned properties did not show specific trends along the vertical direction of the Taiwania stems. [ABSTRACT FROM AUTHOR]

Among the main parameters contributing to the characterization of firewood exploitation modes, the age of the collected wood is very difficult to assess in anthracology. In this paper, we propose an original way to investigate the age of the exploited wood, the heartwood formation process which takes place in 20–25-year-old deciduous oak trees. The formation of tyloses in earlywood vessels is an important feature of the changeover of sapwood to heartwood. However, tylosis formation also occurs in sapwood. Therefore, the observation of the mere presence of tyloses in vessels, as routinely performed in anthracology, is not sufficient to distinguish sapwood from heartwood. The aim of this study was thus to use the proportion of vessels sealed by tyloses as a discriminating marker between sapwood and heartwood applicable to anthracology in order to characterize firewood exploitation in the past. The trunks and branches of ten deciduous oak trees ranging from 15 to 60 years old were sampled in three French sites. For an application to archaeological charcoal (tyloses are preserved until 800 °C), thresholds of less than 65% for sapwood and up to 85% for heartwood appear to be significant at the population level for deciduous oak ( Quercus petraea/robur ) in a temperate climate. Besides the indication of the minimal age ( ca. 25 years old in the case of deciduous oak), the variability of the proportion of vessels with tyloses in sapwood is explored as a good indicator of the vitality of the wood which opens new prospects for the exploitation of dendro-anthracological parameters, such as the discrimination of branch wood. [ABSTRACT FROM AUTHOR]

The norlignan (NorL) content in extractives of sugi (Cryptomeria japonica) shows a decreasing tendency from the outer to the inner part of the heartwood (hW). The hypothesis, that this effect may be due to polymerization, which aggravates NorL extraction, has been explored in the present study. To this purpose, samples were collected and extracted from the outer, middle and inner parts of the hW. Fractionation procedures of NorL-oligomers by LH-20 column chromatography were established, and the fractions were characterized by advanced analytical methods such as HPLC-SEC and HPLC combined with electrospray ionization ion trap mass spectrometry (HPLC-ESI-ion-trap-MS). Agatharesinol and sequirin C were the most abundant compounds. Several individual NorL-dimers and -trimers were detected for the first time, and four kinds of NorL-dimers and seven kinds of NorLtrimers were semi-quantitatively analyzed by HPLC-ESIMS combined with selected ion monitoring (SIM). The oligomer contents show an increasing tendency from the outer part toward the middle and inner part of the hW, which accounts mainly for the monomeric NorL decrement in the same direction. [ABSTRACT FROM AUTHOR]