"IAWA List of Microscopic Bark Features" published on 22 Nov 2016 by Brill.

The specific pattern of the wood of Karelian birch ( Roth var. (Merckl.) Hämet-Ahti), is created mainly by dark-coloured inclusions of parenchyma tissue. Our study revealed that the greatest density of parenchyma inclusions in Karelian birch wood is observed above branch attachments to the trunk and below forks. In the place of branch attachment, phloem flows of photoassimilates (sucrose) from the branch and along the trunk merge into one pathway, causing a rise in sucrose content in tissues there. In the area below the fork, sucrose flows from two (or more) trunk axes are combined. Many studies have demonstrated that elevated sucrose level is associated with the differentiation of parenchyma. We believe that where large phloem fluxes merge a high level of sucrose promotes mass differentiation of parenchyma cells instead of fibers and vessels. As a result, the density of the figured pattern in the wood increases. The obtained data have a practical value and can be used in developing recommendations for Karelian birch cultivation.Betula pendula carelica

Auxin status in woody plants is believed to be a critical factor for the quantity and quality of the wood formed. It has been previously demonstrated that figured wood formation in Karelian birch (Betula pendula Roth var. carelica (Merckl.) Hämet-Ahti) is associated with a reduced auxin level and elevated sugar content in the differentiating xylem, but the molecular mechanisms of the abnormal xylogenesis remained largely unclear. We have identified genes involved in auxin biosynthesis (Yucca), polar auxin transport (PIN) and the conjugation of auxin with amino acids (GH3) and UDP-glucose (UGT84B1) in the B. pendula genome, and analysed their expression in trunk tissues of trees differing in wood structure. Almost all the investigated genes were overexpressed in Karelian birch trunks. Although Yucca genes were overexpressed, trunk tissues in areas developing figured grain had traits of an auxin-deficient phenotype. Overexpression of GH3s and UGT84B1 appears to have a greater effect on figured wood formation. Analysis of promoters of the differentially expressed genes revealed a large number of binding sites with various transcription factors associated with auxin and sugar signalling. These data agree with the hypothesis that anomalous figured wood formation in Karelian birch may be associated with the sugar induction of auxin conjugation.

The mechanisms regulating the tree trunk radial growth can be studied in original experiments. One technique for studying cambium activity (the meristem involved in radial growth) under conditions of an increased photoassimilate level is trunk girdling. We girdled the trunks of 17- to 22-year-old silver birch plants (Betula pendula Roth var. pendula) during the active growth period and collected xylem and phloem samples at two height levels (1 cm and 35 cm) above girdle, 10, 20, and 30 days after girdling. We investigated the changes that occurred at the anatomical level, as well as the activities of sucrose-metabolizing enzymes and antioxidant-system enzymes and the expression of genes that encode proteins involved in sucrose and auxin transport and metabolism. A moderate increase in photoassimilates (35 cm above the girdle) resulted in a change in the ratio of phloem to xylem increments and an increase in the proportion of parenchyma in the conducting tissues. The increase of photoassimilates above the level at which they can be used in the processes of normal tissue growth and development (1 cm above the girdle) led to xylogenesis suppression and the stimulation of phloem formation, a significant increase in the parenchyma proportion in the conducting tissues, and formation of large sclereid complexes. The differentiation of parenchyma and sclereid cells coincided with biochemical and molecular markers of abnormal conducting tissue formation in Karelian birch, which are also characterized by high proportions of parenchyma and sclereid near the cambium. The results obtained are important in understanding the cambium responses to the photoassimilate distribution changes and estimating tree productivity and survival under changing environmental conditions.

This study is a continuation of research on the role of sucrose in figured wood formation in temperate trees. Different concentrations of sucrose solutions were administered for 7 weeks to trunk tissues ofBetula pendulaRoth, Alnus incana(L.) Moench andPopulus tremulaL. Then xylem anatomy was examined with particular emphasis to the number of vessels and the spatial orientation of xylem elements. InB. pendulaandA. incanaa high level of exogenous sucrose caused a reduction in the number and size of xylem vessels, even to the point of absence of vessels. Sucrose concentrations of 100 and 200 g l^-1induced the formation of curly grain and anomalous club-shaped rays in xylem ofB. pendula.Populus tremulaxylem was not significantly altered by the experiment; the xylem anatomy was more seriously affected by wounding than by sucrose. InB. pendulaandA. incanathe wood formed during the experiment was similar to figured wood of these species. The decrease in the number and size of vessels in the xylem formed during the experiment possibly suggests that high concentrations of sucrose lead to a decline in the level of physiologically active auxin. Changes in the orientation of xylem elements points to a disruption of basipetal auxin transport. Further biochemical and physiological studies are needed to provide more comprehensive understanding of the relationship between sucrose and auxin during the development of figure in wood.