Comparative analysis of diterpene composition in the bark of the hybrid larch F<Subscript>1</Subscript>, <Emphasis Type="Italic">Larix gmelinii</Emphasis> var. <Emphasis Type="Italic">japonica × L. kaempferi</Emphasis> and their parent trees

The diterpene compositions in the bark of branches were investigated for two families of the F₁ hybrid, Kurile larch (Larix gmelinii var. japonica Pilg.) × Japanese larch [Larix kaempferi (Lamb.) Carr.] (hereafter F₁) and their parents clones. 13-Epimanool, larixol, larixyl acetate, 13-epitorulosyl acetate (not detected in L. gmelinii var. japonica), isopimaric acid, abietic acid, dehydroabietic acid, and neoabietic acid were detected. Larixol and abietic acid represented more than 50% of the diterpene content in L. gmelinii var. japonica and L. kaempferi, respectively. Larixol and abietic acid were the predominant diterpene components in the F₁, and the proportions of these diterpenes were between those of the parental species. Therefore, the diterpene compositions in the F₁ were hereditarily infl uenced by their parents. The ratios of labdane, pimarane, and abietane diterpenes suggested that the main diterpene biosynthesis pathway in L. gmelinii var. japonica was from copalyl diphosphate (CDP) to labdane-type diterpenes, and that in L. kaempferi was from CDP to abietane-type diterpenes via pimarane type. Furthermore, linear discriminant analysis suggested that the diterpene contents are effective indices for the discrimination of the hybrid seedlings. Part of this article was presented at the Annual Meetings of the Japan Society for Bioscience, Biotechnology, and Agrochemistry in Kyoto, March 2006, and in Tokyo, March 2007     

Characteristics of reproductive biology forLarix originating in Japan

The paper summarized the life cycle, environmental features and distributions ofLarix origination in Japan. The time of blooming and fruiting and the ways of cone collection and seeds storage were introduced. The treating methods of seeds germination and the sowing time for both Japanese larch (Larix kaempferi Carr) and Dahurian larch (Larix gmelinii var.japonica Pilger) were discussed. Responsible Editor: Zhu Hong     

The effects of early and intense pruning on light penetration,tree growth,and epicormic shoot dynamics in a young hybrid larch stand

We examined the effect of early and intense pruning on light intensity under the canopy, individual growth, diameter–height relationships, and epicormic shoot dynamics in young hybrid larch (Larix gmelinii var. japonica × L. kaempferi) to establish a new effective management method for hybrid larch plantations. The objective is to produce high-quality wood while reducing silviculture costs using a combination of low-density planting and early and intense pruning. In a young hybrid larch plantation, we pruned branches to two different heights (2 and 4 m above ground level) using a no-pruning treatment as a control. Although the growth rates were lower in the heavy pruning treatment (4 m above the ground level) than in other treatments in the year following pruning, when measured 4 years later, growth did not differ between treatments. The number of epicormic shoots increased in the year following pruning, as did the relative photosynthetic photon flux density (rPPFD). The number of epicormic shoots was also dependent on the size of individual trees. However, survival of epicormic shoots was not sufficiently high to be problematic for high-quality timber production. If branches are pruned carefully such that the rPPFD does not rise above 20%, the emergence of epicormic shoots can also be controlled. Our results indicate that early and intense pruning is an effective component of a new management system for hybrid larch plantations.     

Activities of some defense proteins associated with age or plant family in larch needles of <Emphasis Type="Italic">Larix olgensis</Emphasis> and <Emphasis Type="Italic">L. kaempferi</Emphasis> × <Emphasis Type="Italic">L. gmelinii</Emphasis>

Defence proteins are a kind of chemical defence compounds. They play a key role in plant restraining biotic and abiotic harm. To illuminate activities of some defense proteins associated with age or plant family of larch, the larch needles were collected from two different families of Korean larch, Larix olgensis and a hybrid larch, L. kaempferi × L. gmelinii, respectively, and then the activities of defence proteins in those were tested using a UV spectrophotometry. The results showed that the activities of protective enzymes at the16-/17- and 19-year-age groups were higher than those at the other age groups in the both larch species. While the activities of polyphenol oxidase (PPO) at the 16-/17-year-age group and phenylalanine ammonia-lyase (PAL) and protease inhibitors at the 19-year-age group were the highest among all age groups. Then, compared with L. olgensis, the hybrid larch had significant effects on the activities of protective enzymes. The effects of plant family on the activities of PAL and chymotrypsin inhibitor were significantly different, and then those on the activities of PPO were not significantly different. The effects of the two families in L. olgensis on the activities of trypsin inhibitor (TI) were significantly different, while those in the hybrid larch on the activities of TI were contrary. To conclusions showed that the different age and plant family significantly affected the activities of defence proteins in the needles of two larch species, and then enhanced the larch resistance to pests. These could play a key function in forestry tree genetic improvement and management in future.     

Age trends in the genetic parameters of wood density and the relationship with growth rates in hybrid larch (Larix gmelinii var. japonica × L. kaempferi) F1

Age trends in variance components and heritability of overall wood density, earlywood and latewood density, and latewood proportion were investigated in 29-year-old trees of 19 full-sib families of hybrid larch (Larix gmelinii var. japonica × Larix kaempferi) F₁. The age–age correlation and optimum selection age for these traits were also estimated and genetic and phenotypic correlations between wood density and radial growth rate were calculated for each growth ring. Intraring wood density data were obtained using X-ray densitometry. The coefficient of additive genetic variance was stable over all ages, whereas the coefficient of environmental variances gradually decreased with increasing age, resulting in increases in heritability estimates with age for overall density. The latewood proportion had the highest heritability estimates at all ages, ranging from 0.44 to 0.66. Overall density and its various components at 28 years of age showed strong genetic correlations with their respective traits at all younger ages. Optimum selection ages for the wood density traits ranged from 8 to 14 years, at which point maximum gain efficiencies per year were obtained. There were negative correlations between wood density and radial growth rate at early ages, although these relationships tended to be weaker with increasing age. These results suggest that selection at a young age is effective for wood density, but particular care must be taken in selecting trees with an improved radial growth rate because rapid growth will result in a low-density wood product, especially in the early growth period.     

Genetic control of intra-ring wood density variation in hybrid larch (Larix gmelinii var. japonica × L. kaempferi) F1

Genetic parameters for various wood density traits were estimated in 29-year-old trees of 18 full-sib families of hybrid larch (Larix gmelinii var. japonica × Larix kaempferi) F₁. Intra-ring density variation (IDV) was also evaluated using a model that expresses the pattern curve from earlywood to latewood as a power function. A high IDV indicates an abrupt change in wood density from earlywood to latewood. The ring width and wood density traits of individual rings were determined by X-ray densitometry. Overall wood density (RD) was shown to increase with increasing ring number, ranging from 0.42–0.59 g/cm³, whereas IDV of individual rings decreased gradually from pith outwards. Estimates of individual tree narrow-sense heritability of RD and IDV were 0.66 and 0.67, respectively. IDV showed negative genetic and phenotypic correlations with RD (r _g = −0.99, r _p = −0.72). The predicted genetic gains in latewood proportion and IDV were higher than that of RD. These results suggest that the intra-ring density variation is under moderate genetic control equivalent to wood density. The trend of increasing wood density from earlywood to latewood was associated with changes in both tracheid diameter and cell wall thickness.     

落叶松种间及其杂种生长与形质性状评价研究

[目的]通过对落叶松种间及其杂种生长与形质性状评价来定向选育优良落叶松建筑材。[方法]以辽宁省大孤家林场32年生的落叶松试验林为实验材料,对落叶松属日本、兴安、长白及华北4个落叶松种,及以日本落叶松为母本的日本×兴安、日本×华北、日本×长白、日本×日本杂种的生长和形质性状进行变异规律分析,并采用模糊数学隶属函数对其差异显著指标进行综合评价。[结果]表明:落叶松种和杂种间生长差异显著(p<0.01),杂种胸径、树高和材积总生长量与母本差异不显著,却显著高于相应父本;杂种之间的生长差异体现在林分的前10年,随着林龄的增大优势逐渐减退;树冠特征因子中的枝下高、冠长、枝条直径、长度、角度和干形特征因子中的高径比、树干圆满度、节子直径、长度、角度在落叶松种和杂种间差异显著。基于上述指标进行了生长和形质性状综合评价,排名前三位的分别为日本×兴安、日本×长白和日本落叶松。32年生时日本×兴安的胸径、树高和材积分别为18.69 cm、21.62 m和0.339 2 m³;日本×长白的为18.44 cm、23.85 m和0.339 6 m³;日本落叶松的为19.36 cm、21.31 m和0.335 7 m³;而且它们在适应性上也具有明显优势。[结论]日本×兴安、日本×长白和日本落叶松适合作为定向培育建筑材的优良品种在东北地区重点推广。     

Resistance of larches to bark stripping by sika deer in Hokkaido,northern Japan

Resistance of the Japanese larch (Larix kaempferi) (Lk), the Kurile larch (L. gmelinii var. japonica) (Lg), and their F₁ hybrid (Lg × Lk) to bark stripping by sika deer (Cervus nippon yesoensis) were studied in a forest in Hokkaido by cafeteria tests. Several sets of logs prepared from Lk, Lg, and Lg × Lk were fed to the deer. The stripped areas of logs significantly differed between the Lk/Lg set and the Lk/Lg × Lk set (P 0.05). The resistances of Lg and Lg × Lk to bark stripping was stronger than Lk. This result was similar to those of previous studies on gray-sided voles and mountain hares in Hokkaido.     

Estimated amount of carbon accumulation of hybrid larch in three 31-year-old progeny test plantations

Hybrids generated by crossing Kuril larch (Larix gmelinii var. japonica) and Japanese larch (L. kaempferi) are expected to have high carbon accumulation ability because of fast growth and high wood density in Hokkaido, Japan. We estimated the amount of carbon accumulation of the hybrid larch in three progeny test plantations consisting of 21 full-sib families and compared the results to the carbon accumulation of open-pollinated progenies of Japanese larch plus-trees (improved Japanese larch). Gene-environment interactions were not observed for tree height and diameter at breast height but were seen for wood density (area-weighted density, AWD). The amount of carbon accumulated per unit area (C_stand) positively correlated with the stand volume. The AWD did not correlate with the C _stand; therefore, families with high wood density can be selected independent of the C _stand. The C _stand of the best full-sib family, female parent half-sib family, and male parent half-sib family at three sites were 106.1, 84.6, and 93.2 Cton·ha⁻¹, respectively. All these values exceed the mean for the improved Japanese larch, which has a C _stand value of 82.5 Cton·ha⁻¹.     

Age trends of genetic parameters of spiral grain in hybrid larch F1 and implications for efficiency of early selection

Age trends in variance components and heritability were estimated from the spiral grain angle of rings 2–25, as counted from the pith in hybrid larch (Larix gmelinii var. japonica × Larix kaempferi) F₁. Age-age genetic and phenotypic correlations and the optimum selection age for spiral grain were also calculated. Wood samples were collected from 95 29-year-old trees belonging to 19 full-sib families in a progeny test plantation in Hokkaido, northern Japan. Spiral grain angle data were obtained by the splitting method. Mean grain angles at growth rings used for the analyses were calculated as the arithmetic mean of angles up to respective rings. Generally, the additive genetic variance for mean grain angle decreased with increasing ring number. Although dominance variance was comparatively high near the pith, it decreased to zero in subsequent growth rings. Highest heritability estimates of mean grain angle occurred at ring 4 and then declined with age, ranging from 0.45 down to 0.20. Age-age phenotypic correlations were higher than genetic correlations, especially those involving early growth rings. Optimum selection ages for spiral grain based on genetic and phenotypic correlations were estimated as 3 and 4 years (cambial age), respectively, in which maximum gain efficiency per year were obtained.     

Genetic variation in wood stiffness and strength properties of hybrid larch (Larix gmelinii var. japonica × L. kaempferi)

Genetic parameters for wood stiffness and strength properties were estimated in a 29-year-old hybrid larch stand (Larix gmelinii var. japonica × Larix kaempferi). The study included 19 full-sib larch families from Hokkaido, northern Japan. Implications of these genetic parameters in wood quality improvement are subsequently discussed. Traits included in the analyses were the dynamic modulus of elasticity of green logs (E _log), the modulus of elasticity (MOE), the modulus of rupture (MOR), compression strength parallel to the grain (CS) in small clear specimens, wood density (DEN), and diameter at breast height (DBH). DEN had the lowest coefficients of variation and MOE the highest. The narrow-sense heritability estimates of E _log, MOE, MOR, and CS were 0.61, 0.44, 0.60, and 0.43, respectively, and those of DEN and all mechanical properties increased from an inner to outer position within the stem. E _log and DEN had high positive phenotypic (0.52–0.83) and genetic (0.70–0.92) correlations with MOE, MOR, and CS. The mechanical properties of the inner position of the stem had rather high phenotypic and genetic correlations with those of the outer position and overall mean. The predicted gains in wood stiffness (E _log and MOE) were higher than those of the strength properties (MOR and CS). The predicted correlated responses in MOE, MOR, and CS when selecting for E _log and DEN were 72.6%–97.8% of a gain achievable from direct selection of these traits. DBH showed an insignificant correlation with all mechanical properties, although selection of this trait had a slightly negative effect on the mechanical properties.     

3种落叶松苗期光能利用效率的比较

正林木90%以上的干物质来自光合作用,单位时间内光能利用效率(包括光能截获能力和光能转化效率)是衡量林木生长潜力的重要指标。光能截获能力主要与叶面积大小及其发展有关,光能转化效率主要与单叶光合速率有关。自发现光合速率在不同个体间存在明显差异后,以提高光能利用效率为目标的"高光效育种"受到育种学家的重视[1-3],而通过选配高光效种质进行有目的杂交已成为高光效育种的重要手段[4]。落叶松(Larix spp.)种间杂交     

Eight-year performance of bareroot Douglas-fir and bareroot and plug western larch Seedlings following herbicide applications,northeast Oregon,USA

Plantation success in northeastern Oregon can be limited by poor survival for western larch (Larix occidentalis Nutt.) and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. glauca (Beissn.) Franco) seedlings. Vegetation management treatments may increase soil moisture availability during the dry summer months, thereby increasing the probability of seedling survival. Chemical site preparation treatments were applied in spring 2007, summer 2007, and spring 2008 to determine if such treatments would improve survival and growth of bareroot Douglas-fir and bareroot and plug western larch seedlings on three sites near Elgin, Oregon. Although some herbicide treatments significantly increased survival, results were not consistent among sites. Overall, Douglas-fir had greater survival after 7–8 years than larch seedlings. Seedling size 7–8 years after planting was also highly variable among sites and treatments with larch seedlings being taller and larger than Douglas-fir seedlings, on the average. Size for all stock types was negatively correlated with vegetative cover and positively correlated with first-year soil moisture. Macro- and microclimatic conditions, including topography and soil depth, were likely contributors to differences in survival and growth.     

Effects of silviculture treatments in a hurricane-damaged forest on carbon storage and emissions in central Hokkaido,Japan

Hurricanes cause abrupt carbon reduction in forests, but silviculture treatment can be an effective means of quickly regenerating and restoring hurricane-damaged sites. This study assessed how silviculture treatments affect carbon balance after hurricane damage in central Hokkaido, Japan. We examined carbon storage in trees and underground vegetation as well as carbon emissions from silviculture operations in 25-year-old stands, where scarification and plantation occurred just after hurricane damage. The amount of carbon stored varied according to silviculture treatment. Among three scarification treatments, a scarified depth of 0 cm (understory vegetation removal) led to the largest amount of carbon stored (64.7 t·ha-1 C). Among four plantation treatments, the largest amount of carbon was stored in a Larix hybrid (L. gmelinii var. japonica × L. kaempferi) plantation (80.3 t·ha-1 C). The plantation of Abies sachalinensis was not successful at accumulating carbon (40.5·ha-1 C). The amount of carbon emitted from silviculture operations was 0.05-0.14 t·ha-1 C, and it marginally affected the net carbon balance of the silviculture project. Results indicate that silviculture treatments should beperformed in an appropriate way to effectively recover the ability of carbon sequestration in hurricane-damaged forests.     

Early evaluation of growth traits of <Emphasis Type="Italic">Larix kaempferi</Emphasis> clones

Early selection is an important method to shorten the breeding cycle for tree species, which may differ in the time for early selection. To evaluate the early selected time for Larix kaempferi, tree height and diameter at breast height of 57 L. kaempferi clones were measured over many different growth years. The results indicated that, except for age × clone interaction for diameter at breast height (P = 0.741), there were significant differences among all variation sources (P < 0.01). The coefficient of phenotypic variation ranged from 14.89 to 35.65% for height and from 19.17 to 23.86% for diameter at breast height in different growth years. The repeatability of height and of diameter at breast height among clones was high, ranging from 0.6181 to 0.8531 (height) and from 0.8443 to 0.8497 (diameter at breast height), in different growth years. There were significant positive correlations between all pairs of growth traits except between height in the 2nd growth year and height in the 30th growth year; and between height in the 2nd growth year × diameter at breast height in the 30th growth year. With a comprehensive evaluation method and a selection ratio of 10%, L65, L1, L62, L9, L15, and L78 were selected as excellent clones in the 30th growth year. Their average values of height and diameter at breast height were 9.81 and 16.57% higher than the overall average, representing genetic gains of 6.46 and 13.99%, respectively. This study provides a theoretical foundation for the genetic improvement of L. kaempferi.     

Distribution of heartwood extractives in hybrid larches and in their related European and Japanese larch parents: relationship with wood colour parameters

Heartwood is the most valuable part of larch timber when either natural durability or aesthetic aspects of the wood are required. Both properties are directly linked to chemical extractives and particularly to phenols. Based on a broad sample of trees (583) from European and Japanese larch and their interspecific hybrid, we investigated the variability in phenolic content, and particularly of two major compounds, Taxifolin (Tax) and Dihydrokaempferol (DHK), and their link with wood colour. At the individual wood sample level, phenolic contents ranged from 6.0 to 55.9 mg eq. Tax/g DW. Taxifolin was the most abundant constituent (range: 1.3–41.7 mg/g DW) compared with DHK (0.5–13.7 mg eq. Tax/g DW). A high variability among taxa, genotypes and individual trees within taxa and within trees was observed. Japanese larch had the highest amount of total phenols and of Taxifolin and European larch the lowest. For DHK, Japanese larch was the poorest compared with European larch. Hybrid larch had both a high content of Taxifolin and of DHK. Variability for colour parameters was on average weaker than for phenolic content but still large enough to show significant differences between taxa. Correlations between colour parameters and extractives were moderate to weak. At the mean genotype level, a good link (r > 0.51, p < 0.001) was found between a* (red–green axis in CIELAB) and total phenols and Taxifolin contents and between L* (Lightness) and DHK content (r > 0.54). The broad variability observed in this study at different levels for phenolics offers opportunities for breeders to genetically improve the quality of larch heartwood, in particular in relation to decay resistance.     

Classification of the hot air heat treatment degree of larch wood using a multivariate analysis of near-infrared spectroscopy

In this study, hot air heat treatments of larch (Larix kaempferi) wood specimens were conducted at various temperatures (160–220 °C) and times (1–12 h) to classify the degree of hot air heat treatment using near-infrared (NIR) spectroscopy. NIR reflectance spectra were acquired from specimen cross-sections and were then preprocessed using the standard normal variate. Hierarchical clustering analysis (HCA) using the complete linkage and squared Euclidean distance was conducted to classify the three degrees of heat treatment. A principal component score plot of the NIR spectra was well grouped by the HCA grouping result, and the first component reflected the cluster analysis grouping well. A partial least squares discriminant analysis was performed to develop the discriminant regression model of the three heat treatment degrees. The R² and root mean square error of validation were 0.959 and 0.191, respectively. NIR is considered to be a good candidate to routinely measure the degree of hot air treatment for larch wood.     

Accumulation of constitutive diterpenoids in the rhytidome and secondary phloem of the branch bark of Larix gmelinii var. japonica

The quantitative compositions of the major constitutive diterpenoids in the rhytidome and secondary phloem of the branch bark of Kuril larch (Larix gmelinii var. japonica) were investigated. The eight major diterpenoids were isolated from a diethyl ether extract of the branch bark of L. gmelinii var. japonica and identified as 13-epimanool (1), larixol (2), larixyl acetate (3), 13-epitorulosyl acetate (4), abietic acid (5), neoabietic acid (6), dehydroabietic acid (7), and isopimaric acid (8). The amount of each diterpenoid was subsequently quantified in both the rhytidome and secondary phloem. All of the diterpenoids were present in both bark tissues, but the amounts were significantly higher in the rhytidome than in the secondary phloem. Developed fusiform resin cavities containing oleoresins were commonly observed in dead secondary phloem captured into the rhytidome of a bark transverse section. The accumulation and distribution of these constitutive diterpenoids in the bark tissues can probably be attributed to terpenoid biosynthesis in the living secondary phloem and the rhytidome formation process. From the viewpoint of constitutive chemical defense in conifers, it is suggested that the tree body may be more effectively defended against natural enemies by the higher amount of diterpenoid amount in the outermost and dead bark tissue, the rhytidome, where the potentially poisonous and easily oxidizable diterpenoids can be safely and stably maintained.     

Fall fertilization enhanced nitrogen storage and translocation in Larix olgensis seedlings

Fall nutrient loading of deciduous forest nursery seedlings is of special interest because of foliage abscission and varied translocation patterns. For non-deciduous seedlings in the nursery, fall fertilization typically can reverse nutrient dilution and possibly increase nutrient reserves; however, this technique has received little attention with deciduous conifer trees that translocate nutrients before abscising foliage. We evaluated how fall nitrogen (N) fertilization affected N storage and translocation in the deciduous conifer Olga Bay larch (Larix olgensis Henry) seedlings during the hardening period. Seedlings were supplied with 25 mg N seedling^?1 for 15 weeks before hardening and fall fertilization treatments began with a three week application period of K¹⁵NO₃ at 0, 5, 10 and 15 mg N seedling^?1. During the hardening period, fall N fertilization had little effect on seedling morphology. The N concentration and content of needles decreased dramatically as needles abscised, while that of stems and roots increased. Six weeks after fall N fertilization ceased, all seedlings translocated similar net N from their needles. For the control seedlings, this accounted for 84 % of the N stored in stems and roots. For fall fertilized seedlings, however, the proportion of N stored in stems and roots translocated from needles accounted for only 41–61 % of the total because of absorption of fall fertilizer that was translocated directly to stems and roots. Six weeks after fall fertilization, the distribution pattern of N concentration and content in seedlings was found in this order: stems > fine roots > coarse roots > needles. Our results suggest that providing deciduous conifer seedlings N during hardening, in this case Olga Bay larch, is a way to promote nutrient loading during nursery production.     

Performance of Larix sibirica and L. gmelinii in north-central North Dakota

Siberian larch (Larix sibirica Ledeb.) is a potential alternative tall-tree species for use in shelterbelt plantings in the northern Great Plains. Information from a 26-year-old study containing six Larix seed sources indicates that L. sibirica generally outperformed L. gmelinii and putative L. sibirica × L. gmelinii hybrids in block plantings at a single site in north-central North Dakota. These results suggest that L. sibirica merits further study as a possible species for use in shelterbelt plantings.