Scots pine (Pinus sylvestris L.) wood is desired in woodworking industries due to its favorable timber characteristics and natural durability that is contributed by heartwood extractives. It has been discussed whether the Scots pine heartwood extractives (mainly stilbenes and resin acids) are synthesized in the cells of the transition zone between sapwood and heartwood, or if they are transported from the sapwood. Timing of heartwood formation during the yearly cycle has also not been unambiguously defined. We measured steady-state mRNA levels in Scots pine transition zone and sapwood using RNA sequencing. Year-round expression profiles of selected transcripts were further investigated by quantitative RT-PCR. Differentially accumulating transcripts suggest that, of the Scots pine heartwood extractives, stilbenes are synthesized in situ in the transition zone and gain their carbon-skeletons from Suc and triglycerides. Resin acids, on the other hand, are synthesized early in the spring mainly in the sapwood, meaning that they must be transported to the heartwood transition zone. Heartwood formation is marked by programmed cell death that occurs during the summer months in the transition zone.
To exploit the variation in the natural durability of heartwood timber, screening of a large number of trees or timber is necessary. We suggest that the concentration of total phenolics, measured by the Folin–Ciocalteu (FC) assay, can be used to supplement or even replace in vitro decay tests for screening the variation in the resistance of Scots pine (Pinus sylvestris L.) heartwood timber against the brown-rot fungus Coniophora puteana (Schum. ex Fr.). We screened the juvenile heartwood of 520 Scots pine trees with the FC assay. Samples from 40 trees, with total phenolics concentrations ranging from 1.9 to 21.7 mg tannic acid equivalents (TAE)/g of heartwood, were subjected to an in vitro decay test. The correlation (r) between the total phenolics concentration and mass loss was –0.82 (p < 0.001) in the 7 week long decay test. The relationship was expressed with a linear regression equation: mass loss (mg/g) = 487 – 25 × concentration of total phenolics (mg TAE/g). This strong relationship could be exploited by replacing the decay test with the FC assay to more rapidly assess the durability of wood products.
In this greenhouse experiment, 3-year-old Scots pine (Pinus sylvestris L.) seedlings were wounded by drilling holes through the stem. In the xylem next to the wound, the concentration of resin acids (RAC) increased, and the production of extractives typical for heartwood (stilbenes) and knotwood (stilbenes and lignans) of mature trees was induced. The induced stilbenes were pinosylvin (PS) and pinosylvin monomethyl ether (PSM), and the lignans nortrachelogenin (NTG) and matairesinol (MR). There was positive phenotypic correlation between concentrations of the different extractives. Except for the RAC, the extractive concentrations showed no correlation with the size of the seedlings. The treated seedlings belonged to half-sib families, which enabled the estimation of the genetic parameters for the response variables. The proportion of heritable variation (heritability, h(2)) in the concentration of PS, NTG and MR varied between 0.71 and 1.03, whereas for PSM and RAC the heritability was lower (0.35 and 0.31). Genetic correlation was significant between PS and PSM (r = 0.55, P = 0.018), and between NTG and MR (r = 0.50, P = 0.033). Heritabilities were also estimated on the basis of the regression of the offspring on their mothers h(2)(0P). These estimates were assessed for the concentration of PS, PSM and RAC in the wound response area of the seedlings and correspondingly in the heartwood of their mothers. The heritability was highest for the concentration of PS h(2)(0P). The findings of this study support the suggestion that the wounding of Scots pine seedlings may facilitate the development of an early testing method for breeding heartwood durability.
Stilbenes are abundant in the heartwood of Scots pine (Pinus sylvestris L.) and are known to have strong antifungal efficiency. In this study, Scots pine sapwood blocks were impregnated with crude heartwood extract containing the stilbenes pinosylvin (PS) and the pinosylvin monomethyl ether (PSM). Impregnated blocks were submitted to brown-rot fungi, Coniophora puteana, Gloeophyllum trabeum, and Rhodonia (Poria) placenta, and fungal growth test and decay test were performed. Both tests showed that the impregnation with a high concentration of stilbenes (60 mg g-1 dry wood) significantly suppressed the growth of fungi and slowed down the decay process of wood blocks, especially in case of G. trabeum. However, chemical analysis showed that PS and PSM were degraded by all the three types of fungi, obviously via modification to resveratrol and methylresveratrol. Rhodonia placenta displayed the fastest rate of degradation. Thus, impregnation with biodegradable stilbenes could be a viable alternative for wood preservation only in service situations with low or transient risk of decay.
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