Fate and influence of western red cedar extractives in mechanical pulping

The methanol extractives from western red cedar mechanical pulps were found to be radically different in composition to the extractives obtained from the heartwood. The major heartwood extractive components, the tropolones and lignans, were not present in the extractives from the pulps. However, the proportion of a brown polymer doubled. The low and high molecular weight methanol extractives components from the pulps were separated using methyl tert-butyl ether. The low molecular fraction contained mostly guaiacyl-based compounds with dihydroquercetin, thujic acid, 3-hydroxy-1-(4′-hydroxy-3′-methoxyphenyl)-2-oxopropane and 4-ethyl-2-methoxy-6-hydroxyphenol being positively identified. The brown polymeric portion had molecular weights ranging from 1,000 to 10,000. Infrared analysis indicated that the polymers were formed from lignans. Examination of the changes in diffuse reflectance UV-visible and infra red spectra of the pulps on extraction with methanol, suggested that the colour resides in insoluble polymers formed from plicatic acid/plicatin during refining. Received 18 December 1998     

A capillary gas chromatographic method for thujaplicins in western red cedar extractives

Summary A quantitative method for the analysis of thujaplicins and thujic acid is described. The samples were methylated with diazomethane and analyzed using naphthalene as an internal standard. The procedure was useful in the analysis of western red cedar extractives.     

Leaf longevity of western red cedar (Thuja plicata) increases with depth in the canopy

Leaf longevity determines the annual cost of replacing foliage biomass and influences water and nutrient budgets. Longevity is readily estimated in most evergreen species by annual bud scars along the shoot. However, some species with indeterminate growth do not show these annual markers, making estimation of longevity difficult. One of these species is the widespread and economically valuable western red cedar (Thuja plicata J. Donn ex D. Don), for which no dependable estimates of leaf longevity exist. In this study, we estimated leaf longevity for western red cedar by counting growth rings in shoots at the point of leaf abscission. Estimates were obtained on 26 dominant or codominant trees growing in natural stands in a montane forest in northern Idaho, USA. Leaf longevity averaged 8.9 (SE = 0.2) years, but it strongly increased with depth in the canopy (0.3 year m(-1); mean crown depth was 15 m), increasing from a mean of 6.8 years in the upper third of the canopy to 10.6 years in the lower third. The increase in longevity with depth in the crown is consistent with many reports showing that longevity increases in resource-limiting environments. Longevity did not vary significantly with altitude or solar insolation in these montane forests. Among stand-level variables, longevity was correlated only with leaf area index: it increased slightly in stands with high leaf area indices. This approach to longevity estimation may be useful for any species that produces annual rings but no obvious bud scars, including many Cupressaceae species.     

Nutrition and bud removal affect biomass and nutrient allocation in Douglas-fir and western red cedar

Seedlings of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and western red cedar (Thuja plicata J. Donn ex D. Don) were grown at high (250 mg l(-1)) and low (20 mg l(-1)) nitrogen (N) supply for a year. Before the second growing season, half of the seedlings in each nutrient treatment were allocated to the other treatment. Half of the seedlings in each nutrient treatment then had all growing points removed. Biomass and N, phosphorus (P) and potassium (K) concentrations of old and new shoots and roots were measured three times in the second year to test the interaction of current-year and previous-year nutrient supply on biomass and nutrient allocation in these two species with different growth habits. Pruned seedlings served as controls. Unpruned seedlings of both species increased in height throughout the second growing season, except for Douglas-fir in the N250 --> N20 treatment. Repeated pruning did not prevent new shoot growth, but resulted in a 12 to 52% reduction in biomass of new shoots and new and old roots. Seedlings receiving a low N supply in the first growing season were more severely affected by pruning than seedings receiving a high N supply. Growth was reduced more by pruning in western red cedar than in Douglas-fir. Concentrations of N, P and K were higher in pruned seedlings than in unpruned seedlings. Although dry weights of all plant parts in all treatments increased throughout the second growing season, some retranslocation of N, P and K was observed from old shoots of both species in the N250 --> N20 and N20 --> N20 treatments after August. Quantities of N, P and K retranslocated were greatest in seedlings grown the previous year in the high-N treatment.     

Radial distribution of thujaplicins in old growth and second growth western red cedar (Thuja plicata Donn)

Summary Radial distribution of thujaplicins in western red cedar (Thuja plicata Donn) trees of varying ages was studied in order to assess relative decay resistance of their wood. Samples were extracted with ethanol: benzene (1: 2), and the extractives were analyzed for thujaplicin codtent by a new method utilizing capillary gas chromatography of their methylated derivatives. The combined concentration of thujic acid and methyl thujate was also determined for each sample. Distribution of extractives, thujaplicins and thujic acid, generally increased from pith to outside heartwood, then decreased in the sapwood. Maximum extractive and thujaplicin contents were also related to tree age. This suggests that products made from the wood of younger trees will be less resistant to decay than similar products made from the wood of old trees.     

Testing and evaluation of natural durability of wood in above ground conditions in Europe – an overview

Natural durability of wood is determined by the European standard EN 252 for specimens in ground contact and EN 113 for basidiomycetes in the laboratory, but no test exists for above ground conditions. For above ground conditions, the European prestandard ENV 12037 and EN 330 are used to determine the durability of treated wood. The most important factors for fungal establishment on the surface and within wood are the moisture content, the surrounding temperature, and the relative humidity. Strength tests are the most sensitive for decay detection, but neither strength tests nor identification of fungi responsible for the decay are included in the standards of above ground durability in field tests. To detect decay, visual examination, pick or splinter tests, and mass loss determination are used. Identifying fungi with traditional methods, e.g., growth on solid medium, is time consuming and complicated. Molecular methods like polymerase chain reaction and sequencing do not require mycological skill for identification to species level, and furthermore the methods do not depend on the subjective judgement like most traditional methods, but are based on the objective information of the target organism (e.g., nucleotide sequences). The next generation of standard field tests will probably consider the drawbacks of standard tests today and be rapid and include both quality tests like molecular identification and nondestructive quantitative tests, e.g., acoustic tests. Laboratory tests can be improved by using fungi identified from field trials and by combining different fungi in the same test and thus simulate degradation in practice.     

Phytophthora species associated with dieback of sweet chestnut in Western Turkey

Sweet chestnut (Castanea sativa) is an important tree species in the Marmara and Aegean regions of Turkey as these two regions produce the great majority of edible nuts, especially those used for marron glacé production. Chestnut forests and orchards in these regions showing severe dieback symptoms not associated with chestnut blight were investigated to determine the role of Phytophthora spp. in the decline syndrome. Soil samples were collected from around 108 symptomatic chestnut trees at 29 sites and Phytophthora spp. isolated using soil baiting technique and selective medium. Species isolated were identified by cultural characteristics and ITS sequencing. Phytophthora x cambivora was the dominant species detected in 13 sites, followed by P. cinnamomi (5 sites), P. plurivora (3 sites) and P. cryptogea (1 site). Phytophthora x cambivora was present in both regions, while P. cinnamomi was found only in the Marmara region in coastal areas around Istanbul. When inoculated at the stem bases of 3‐year‐old chestnut saplings, P. cinnamomi produced significantly longer necrotic lesions (7.8–12.0 cm) than P. x cambivora (2.6–6.3 cm) by 12 days after inoculation. Phytophthora plurivora was the least aggressive species causing only small lesions. Phytophthora cryptogea, which represents the first record on chestnut in Turkey, produced intermediate sized lesions in between P. x cambivora and P. plurivora. These results indicate that P. x cambivora and in some areas P. cinnamomi play major roles in the observed dieback of sweet chestnut in western Turkey.     

Root morphology and nutritional status of Japanese red cedar saplings subjected to in situ levels of aluminum in forest soil solution

There is little information on the effects of in situ levels of aluminum (Al) in the forest soil solution on the root morphology of Japanese red cedar (Cryptomeria japonica D. Don). We evaluated whether morphological and nutritional changes in the white roots of Japanese red cedar saplings grown in glass-bead culture for 18 weeks occurred in response to Al solutions at five concentrations: 0 (control), 0.05, 0.1, 0.5, and 1.0mM. Branching white roots treated with 0.5 or 1.0mM Al had some stunted, brown, thickened tips. Their mean lengths were significantly shorter than those of roots treated with 0.05 or 0.1mM Al. The maximum diameters of white roots treated with 0.5 or 1.0mM Al were significantly larger than those of the control. Treatment with 0.5 or 1.0mM Al reduced the concentrations of Ca and Mg in the white roots and increased the concentration of Al compared with the control. These results indicate that between 0.1 and 0.5mM Al, a drastic change occurs in the effects of Al on the morphology and nutritional status of white roots of Japanese red cedar saplings, and suggest that the Al levels in Japanese forest soils may induce morphological changes in the white roots.     

Diversity of Ophiostomatales species associated with conifer-infesting beetles in the Western Carpathians

Bark beetles (Coleoptera: Curculionidae, Scolytinae) are commonly recognised as important agents of tree mortality in coniferous forests of the Western Carpathians. They, together with weevils, are consistently associated with ophiostomatoid fungi. Information regarding conifer beetle-associated fungi in the Western Carpathians remains incomplete and unreliable, particularly with respect to fir-infesting bark beetles. This study aims to clarify associations between fungi in the genera Graphilbum, Leptographium, Ophiostoma and Sporothrix (Ophiostomatales) and their beetle vectors in Norway spruce (Picea abies), European larch (Larix decidua) and silver fir (Abies alba). Samples associated with 20 bark beetle species and weevils were collected from nine stands in Poland and the Czech Republic. Fungi were isolated from adult beetles and galleries. Isolates were identified based on morphology, DNA sequence comparisons for four gene regions (ITS, LSU, ß-tubulin, TEF 1-α) and phylogenetic analyses. In total, 46 distinct taxa were identified, including 25 known and 21 currently unknown species. Several associations between fungi and subcortical insects were recorded for the first time. In addition, O. borealis and O. quercus were detected from A. alba for the first time. The composition of the fungal communities varied among the studied tree species and to a lesser degree among the beetle species. The spruce-infesting bark beetles were commonly associated with species of Leptographium s. l. and Ophiostoma s. str.; the larch-infesting bark beetles were often associated with Ophiostoma s. str. and Sporothrix, while the fir-infesting bark beetles were commonly associated with Ophiostoma s. str. and Graphilbum. The most commonly encountered fungal associates of the examined insects were (a) Grosmannia cucullata, G. piceiperda, Grosmannia sp. 1, Ophiostoma macroclavatum and O. piceae with the spruce-infesting bark beetles; (b) O. pseudocatenulatum and Sporothrix sp. 1 with the larch-infesting bark beetles; and (c) O. piceae, Ophiostoma sp. 2 and Graphilbum sp. 2 with the fir-infesting bark beetles. The differences in fungal associates among the bark beetle species occurring on P. abies, L. decidua and A. alba could be linked to the different habitats that these beetles occupy.     

Decline of radial growth in red oaks is associated with short-term changes in climate

Comparison of basal area increments of paired healthy and declined oak trees shows a marked disjuncture beginning in the early 1950′ at 3 of 4 sample locations across the southeastern United States. An argument is presented that the change in growth was caused (or accelerated) by a series of severe regional droughts in the early 1950′s that impacted the trees which then responded by forming two distinct populations consisting of: 1. relatively healthy trees, and 2. declined trees. Both populations produced less annual basal area increment after several subsequent short-term droughts, but marked crown deterioration and death appeared in the declined population after a moderately severe drought in the early 1980′s.     

Responses of transpiration and photosynthesis to reversible changes in photosynthetic foliage area in western red cedar (Thuja plicata) seedlings

Experiments were conducted on 1-year-old western red cedar (Thuja plicata Donn.) seedlings to determine the response of illuminated foliage to reversible changes in total photosynthetic foliage area (L(A)). Reductions in L(A) were brought about by either shading the lower foliage or by reducing the ambient CO2 concentration (c(a)) of the air surrounding the lower part of the seedling. In the latter case, the vapor pressure was also changed so that transpiration rates (E) could be manipulated independently of photosynthetic rates (A). We hypothesized that following such treatments, short-term compensatory changes would occur in stomatal conductance (g(s)) and A of the remaining foliage. These changes would occur in response to hydraulic signals generated by changes in the water potential gradient rather than changes in the distribution of sources and sinks of carbon within the seedling. When a portion of the foliage was shaded, there was an immediate reduction in whole-seedling E and a concomitant increase in g(s), A and E in the remaining illuminated foliage. However, the intercellular CO2 concentration did not change. These compensatory effects were fully reversed after the shade was removed. When the lower foliage A was reduced to < 0 micromol m-2 s-1, by shading or lowering c(a), and E was either unchanged or increased (by adjusting the vapor pressure deficit), there was no significant increase in g(s) and A in the remaining foliage. We conclude that compensatory responses in illuminated foliage occur only when reductions in L(A) are accompanied by a reduction in whole-plant E. The relationship between the reduction in whole-seedling E and the increase in A is highly linear (r2 = 0.68) and confirms our hypothesis of the strong regulation of g(s) by hydraulic signals generated within the seedling. We suggest that the mechanism of the compensatory effects is a combination of both increased CO2 supply, resulting from increased g(s), and a response of the rate of carboxylation, possibly related to the activity of Rubisco.     

Patterns of regeneration and ground vegetation associated with canopy gaps in a managed beechwood in southern England

In autumn 2001, 15 canopy gaps were selected for study in RumerhedgeWood, a semi-natural, mesotrophic beechwood in southern England.The gaps were located in mature, beech-dominated stands, andhad originated from openings created during a thinning in theearly 1980s and wind damage in 1987/1990 and/or the consequentsalvage operations. The extent of each gap and surrounding treeswere mapped. Tree/shrub regeneration, ground vegetation, bareearth, leaf litter and canopy openness (using a canopy-scope)were measured within and around the gaps using a 5 x 5-m gridand placing a 1 x 1-m quadrat at each grid intersection (totalnumber of quadrats = 400). Most of the gaps were <75 m² inarea. The largest was 241 m². They were generally irregularin shape and there was little or no understorey present. Mostsurrounding trees were beech Fagus sylvatica L. Bramble Rubusfruticosus L. formed a moderate to dense ground vegetation belowmost gaps and declined around the edges only once the gap openingwas substantially obscured. Apart from a few larger saplings,most regeneration was small and of beech. Most of the latterappeared to be in their fifth or sixth growing season, were10–35 cm tall, had an erect base and flat top, had increasedby <5 cm in height during 2001 and were not browsed by deer.Their height and growth form was related to (1) their positionwithin gaps, (2) the degree of canopy openness and (3) the coverof ground vegetation. This was translated into the followingzonation—(1) around the centre of larger gaps: canopyopenness increased to >15 per cent; bramble cover was nearcomplete; litter depth was low; many places had no beech seedlings,but some of the few present were among the tallest, most uprightand fastest growing; (2) towards the edges of the large gapsor directly below smaller gaps: canopy openness was about 4–10per cent; bramble cover was slightly less; beech seedlings weremoderately abundant but patchy, generally shorter, more flattopped and slow growing than in the gap centre, albeit somewere still among the tallest, most upright and fastest growing;(3) beyond the edge of the large/medium gaps (with the gap onlypartially visible) or directly below very small gaps: canopyopenness was only about 2–3 per cent; bramble was muchreduced; beech seedlings were at their most abundant but stillpatchy in distribution and even shorter, more flat topped andslower growing than in the above zone; (4) in an outer zonebeyond or almost beyond the sight of the gaps: canopy opennesswas <2 per cent; bramble was weak and sparse; beech seedlingswere mostly at low densities and predominately short, slow growingand flat topped. This ring pattern of beech regeneration appearedto relate mainly to (1) differences in light availability affectingthe survival, growth rate and form of seedlings; (2) competitionfrom bramble and possibly (3) limited dispersal of beech seedinto gaps. Recommendations are given for managers who wish touse natural regeneration to restock beech woodland.     

Oomycota species associated with deciduous and coniferous seedlings in forest tree nurseries of Western Turkey

Occurrence and pathogenicity of Oomycota species causing root rot were investigated in 10 forest tree nurseries in western Turkey. Soil samples (129 in total) taken from the rhizosphere of symptomatic seedlings were baited for Oomycota using young leaves of Quercus suber, Rhododendron simsii and R. ponticum. Oomycota (178 isolates) were obtained by culturing on selective media, and identified using morphological methods followed by PCR and sequencing of the ITS rDNA and cox1 regions. Phytophthora aff. cactorum, P. citricola sensu lato, P. crassamura, P. syringae, Pythium aphanidermatum, Py. intermedium, Py. irregulare, Py. ultimum and Phytopythium vexans were common amongst the isolates. The highest diversity of Oomycota was found in the forest nurseries at Adapazar?‐Hendek and ?zmir‐Torbal?. Pathogenicity tests showed that the isolates caused lesions on a range of host plants. The importance of these nursery infections in transferring potentially damaging oomycete species to Turkish forests is discussed.     

A molecular marker associated with low-temperature induction of dormancy in red osier dogwood (Cornus sericea)

Dormancy induction in temperate deciduous plants is thought to be regulated by short photoperiods, but low temperature has been shown to eliminate the short photoperiod requirement in northern ecotypes. An F2 population (191 plants) red osier dogwood (Cornus sericea L.) derived from a polycross of an F1 population produced from reciprocal crosses of the parental clonal ecotypes, Northwest Territories (NWT, 62 degrees N) and Utah (42 degrees N), was examined to identify molecular markers of temperature-induced endodormancy. Dormancy induction curves were generated for each individual in the F2 population and a standard point prior to vegetative maturity (i-VM) was inferred from the change in slope of the dormancy acquisition curve. Under Saskatoon, Saskatchewan field conditions (52 degrees N), the NWT ecotype entered i-VM on average 5-6 weeks before the Utah ecotype. Two sub-populations of the F2 population were distinguishable based on VM acquisition on exposure to low temperature but not to short photoperiods. A sequence characterized amplified region (SCAR) marker was developed that correctly (> 92%) identified individual plants within the F2 subpopulation that were responsive to low-temperature induction of VM. Timing of bud break was strongly associated with the timing of VM in the geographical ecotypes but not in the F2 population, indicating that these are separate traits under genetic control.     

Light,soil, and seedling characteristics associated with varying levels of competition in a red pine plantation

Red pine (Pinus resinosa Ait.) seedlings growing under differing levels of competition were evaluated during the fifth growing season following planting, and placed into low, moderate, or high tree classes, as a function of levels of competing vegetation. Tree growth, moisture status, and nutrition were monitored over the growing season. Additionally, site characteristics such as soil temperature and moisture, inorganic nitrogen concentration, mineralized soil nitrogen, and light were measured. Red pine seedlings growing under low competition had an absolute volume growth increase of 795% over the seedlings growing under the heaviest competition. The associated relative volume growth increase during the fifth growing season was 44%. Discriminant analysis was used to describe three classes of trees representing low, moderate, and high levels of competition. Trees growing under low competition had longer and heavier needles, but generally lower nutrient concentrations. Pre-dawn plant moisture status did not vary among competition levels. Soil variables indicated that, in general, the trees growing under low competition occupied warmer, drier, and less fertile microsites, as inorganic soil N and mean monthly mineralized NO₃-N and NH₄>-N tended to be lower on these microsites. The suite of independent variables was effective in classifying the model data into three tree competition classes, with percent correct classifications of 92, 75, and 100 for low (tree class one), moderate (tree class two), and heavy (tree class three) competition, respectively.     

Species richness of the understory woody vegetation in Japanese cedar plantations declines with increasing number of rotations

The possibility of restoring natural broadleaf forests may be decreased by the effects of plantation management, particularly in sites that undergo repeated rotation. We investigated the following two working hypotheses about the effects of repeated plantation of conifers on the natural regeneration of woody saplings in cool-temperate Japanese cedar plantations: (1) that repeated plantation of conifers decreases sapling species richness, and (2) that repeated plantation of conifers changes sapling species compositions. Our result supported the first hypothesis, because species richness was significantly lower in second-rotation plantations than in first-rotation plantations. The second hypothesis was not supported, because no significant or substantial differences in species composition were observed between plantations with different numbers of rotations. However, the abundance of tree (nonshrub) and gravity-dispersed species decreased after the second rotation of large saplings, albeit not those of small saplings, suggesting that response to repeated rotation depended on sapling size. Our results suggest that it is important to consider factors affecting the maintenance of a species in the plantations, such as distance from natural forests and seed sources, to minimize the effects of repeated plantation.     

Comparison of organic matter dynamics in soil between Japanese cedar (Cryptomeria japonica) forest and adjacent Japanese red pine (Pinus densiflora) forest established on flatland

In order to clarify the effects of tree species on organic matter dynamics in soil, we investigated the amount of forest floor material, leaf litter decomposition rate, soil chemical characteristics, soil respiration rate and cellulose decomposition rate in a Japanese cedar forest (cedar plot) and an adjacent Japanese red pine forest (pine plot) established on a flatland. The amount of forest floor material in the cedar plot was 34.5 Mg ha⁻¹ which was greater than that in the pine plot. Because the leaf litter decomposition rate was higher in the pine plot than in the cedar plot, it is likely that the difference in the amount of forest floor material between the plots is caused by the difference in the leaf litter decomposition rate. The C concentrations of soil in the cedar plot were 1.2–2.1 times higher than those in the pine plot. Soil pH(H₂O)s in the cedar plot were significantly higher than those in the pine plot. The soil respiration rates and the rates of mineralized C in the cedar plot byin vitro incubation were higher than those in the pine plot. From this result, it is assumed that soil organic matter in the cedar plot was decomposed relatively faster compared with the pine plot. Furthermore, microbial activities, which were reflected as cellulose decomposition rates in the cedar plot, were higher than those in the pine plot. A part of this paper was presented at the 109th Annual Meeting of the Japanese Forestry Society (1998).     

Effects of soil pH gradient caused by stemflow acidification on soil microarthropod community structure in a Japanese red cedar plantation: An evaluation of ecological risk on decomposition

Effect of acidic stemflow input on soil biological properties around cedar tree trunk were examined in a 34 year old Japanese red cedar plantation forest. Average soil pH of soil around cedar trunk (stemflow soil) was 3.88 and that of soil between trees (throughfall soil) was 4.97, and this suggested soil acidification due to proton load via stemflow. Soil microarthropod density in soil was higher as close to tree trunk, whereas microbial biomass of surface soil was reduced ca. 30 % around trunk. Abundance of Collembola and Gamasid mites showed positive correlation to amount of soil organic matter, while Oribatida had strong negative correlation to soil pH.Tectocepheus velatus (Oribatida) was dominant in soils pH under 4.0. Annual decomposition rate of cedar foliage was reduced ca. 16% in stemflow soil compared to throughfall soil. More litter accumulation was observed in stemflow soil, but it was not significant, though low microbial activity and retarded litter decomposition may be responsible to the accumulation of organic matter in stemflow soil. Soil biological community and soil process have been changed due to stemflow, the affected area corresponded to ca. 9.4 % of the forest. Thus, ecological risk of acidification in the sense of decrease in decomposition rate is 1.4%. This study was partly supported by Grant-in-Aid for Scientific Research (A)(1), No.08506001 by The Ministry of Education, Science, Sports, and Culture, Japan, and by the research project “Establishment of a scientific framework for the management of toxicity of chemicals based on environmental risk-benefit analysis” supported by Core Research for Evolutional Science and Technology (CREST).     

Comparisons of Ophiostomatoid fungi associated withTomicus piniperda andT. minor in Japanese red pine

Five species ofOphiostoma, twoLeptographium species and aGraphium species were isolated from two morphologically and ecologically similar bark beetle species,Tomicus piniperda andT. minor, and their infested Japanese red pine (Pinus densiflora) in Yamanashi Prefecture, central Honshu, Japan. An underscribedOphiostoma species andO. minus were isolated mainly fromT. piniperda and its galleries.Ophiostoma canum which was found for the first time in Japan was mainly fromT. minor and its galleries. Specific relationships between the beetles and fungal species are suggested. Contribution No.140, Laboratory of Plant Pathology and Mycology, Institute of Agriculture and Forestry, University of Tsukuba. Part of this study was presented at 108th Annual meeting of Japanese Forestry Society, April 2, 1997, Fukuoka, Japan.