Freezing tolerance of conifer seeds and germinants

Survival after freezing was measured for seeds and germinants of four seedlots each of interior spruce (Picea glauca x engelmannii complex), lodgepole pine (Pinus contorta Dougl. ex Loud.), Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and western red cedar (Thuja plicata Donn ex D. Donn). Effects of eight seed treatments on post-freezing survival of seeds and germinants were tested: dry, imbibed and stratified seed, and seed placed in a growth chamber for 2, 5, 10, 15, 20 or 30 days in a 16-h photoperiod and a 22/17 degrees C thermoperiod. Survival was related to the water content of seeds and germinants, germination rate and seedlot origin. After freezing for 3 h at -196 degrees C, dry seed of most seedlots of interior spruce, Douglas-fir and western red cedar had 84-96% germination, whereas lodgepole pine seedlots had 53-82% germination. Freezing tolerance declined significantly after imbibition in lodgepole pine, Douglas-fir and interior spruce seed (western red cedar was not tested), and mean LT50 of imbibed seed of these species was -30, -24.5 and -20 degrees C, respectively. Freezing tolerance continued to decline to a minimum LT50 of -4 to -7 degrees C after 10 days in a growth chamber for interior spruce, Douglas-fir and lodgepole pine, or after 15 days for western red cedar. Minimum freezing tolerance was reached at the stage of rapid hypocotyl elongation. In all species, a slight increase in freezing tolerance of germinants was observed once cotyledons emerged from the seed coat. The decrease in freezing tolerance during the transition from dry to germinating seed correlated with increases in seed water content. Changes in freezing tolerance between 10 and 30 days in the growth chamber were not correlated with seedling water content. Within a species, seedlots differed significantly in freezing tolerance after 2 or 5 days in the growth chamber. Because all seedlots of interior spruce and lodgepole pine germinated quickly, there was no correlation between seedlot hardiness and rate of germination. Germination rate and freezing tolerance of Douglas-fir and western red cedar seedlots was negatively correlated. There was a significant correlation between LT50 after 10 days in the growth chamber and minimum spring temperature at the location of seedlot origin for interior spruce and three seedlots of western red cedar, but no relationship was apparent for lodgepole pine and Douglas-fir.     

Supercritical fluid impregnation of selected wood species with tebuconazole

The effects of pressure and temperature on supercritical fluid impregnation of tebuconazole were evaluated on Douglas-fir, western red cedar, red alder, white spruce, and white oak. Higher pressure markedly enhanced both the retention and distribution of tebuconazole in these species. When the rate of pressure release was altered at the ends of treatments of Douglas-fir, results varied. Generally, a higher rate of venting increased the steepness of the preservative gradient inward from the surface. Elevated pressures also affected some wood properties. Western red cedar and white spruce showed collapse, while the other three species were free of such defects. Modulus of Elasticity (MOE) and Modulus of Rupture (MOR) tended to decline with higher pressure in western red cedar and white spruce, but the differences were rarely significant. No significant changes in MOE/MOR occurred with the other 3 species. Received 9 November 1998     

Outdoor exposure of untreated Scots pine (Pinus sylvestris L.) and Norway spruce [Picea abies (L.) Karst.] wood samples

Abstract

Untreated Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) samples were exposed above ground in a durability test for 6 years. The samples consisted of three pieces of wood, 22×95×500 mm, screwed together; two pieces lengthwise with a third piece overlapping. Weight was measured, to calculate moisture content (MC), and samples checked regularly for cracks and fungal growth. Parameters investigated were heartwood/sapwood (pine), annual ring orientation (spruce), stand site, annual ring width and density. Stand site, annual ring width and density had no influence on MC or fungal growth for either pine or spruce. Spruce samples with vertical annual rings had fewer cracks than samples with horizontal annual rings. Pine sapwood samples had a high MC and a large amount of rot fungi, while heartwood had a lower MC and no rot. Most spruce samples were similar to pine heartwood, except from a few samples that had high MC and fungal growth. Those were all sawn from the outer part of the log. Therefore, it can be stated that spruce sawn from the inner part has almost the same properties as pine heartwood, while spruce from the outer part of the log has similar properties to pine sapwood.     

Fluorescein diacetate as a viability stain for tree roots and seeds

Fluorescein diacetate (FDA) was tested as a viability stain for roots of green ash as well as for seeds of green ash and 10 other tree species. The viability level indicated by FDA staining of green ash roots agreed well with root growth potential results, bud condition assessment, and foliage browning measurements. In seed viability experiments, the FDA staining intensity of embryos was related to germination in 9 out of 11 species tested using a 30 minute stain incubation period. In the other 2 species, eastern hemlock and Scotch pine, embryo FDA staining intensity and germination were also similar, provided an 18 h stain incubation period was used. When two seedlots of differing viability were tested in each of white spruce, Douglas-fir, and pitch pine, significantly higher germination was reflected in significantly higher embryo FDA staining intensity. In Sitka spruce seed that was heat treated to produce a range of viabilities, the semilog plot of germination (log scale) and FDA staining intensity of the embryo (linear scale) had an r²= 0.95. Based on these preliminary results, FDA shows promise as a rapid viability stain for tree roots and seeds.     

The effects of Douglas-fir on tree-specific arthropod communities in mixed species stands with European beech and Norway spruce

The ecological effects of planting exotic Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] in Central Europe are still poorly understood. The aim of this study was to answer the question of whether Douglas-fir affects tree specific arthropod communities in different mature forest types (Douglas-fir, spruce and beech dominated) in Southern Germany. Therefore, arthropod communities of stem and tree crown strata of Douglas-fir and spruce (Picea abies L.) were sampled in the years 1999–2001 using arboreal photo-eclectors and flight interception traps. Statistical analysis was conducted for all species and focused on conifer specialists at three levels: (1) species diversity, (2) guild structure and (3) community structure. Within the stem stratum, species diversity was significantly higher on spruce than on Douglas-fir independent of year and stand composition. This could not be explained by a single feeding guild, rather by species changing strata during the vegetation period. In contrast, species diversity in tree crowns was approximately the same for both conifer species. However, communities in Douglas-fir crowns were conspicuously different from those in spruce crowns, especially in the Douglas-fir dominated stand type. While zoophagous insects exhibited higher activity on Douglas-fir in 2000, xylophagous beetles were more abundant on spruce in 2001. In European beech stands with widely spaced Douglas-fir trees, the site specific and broad-leaved tree related fauna might be maintained. In addition, Douglas-fir with its resource of Adelges cooleyi and crowns that overtop the broad-leaved tree canopy, offer additional resources for several aphidophagous and thermophile species.     

Radial patterns of carbon isotopes in the xylem extractives and cellulose of Douglas-fir

Heartwood extractives (nonstructural wood components) are believed to be formed from a combination of compounds present in the adjacent sapwood and materials imported from the phloem. The roles of local compounds and imported material in heartwood formation could have important implications for the wood quality of species having naturally durable wood. Stable isotope composition (delta(13)C) was analyzed to assess radial variation in sapwood extractives, and to estimate the relative importance of adjacent sapwood extractives and imported photosynthate in the formation of heartwood extractives. Cellulose and extractives from the outer 39 annual rings of six Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees were isolated and their delta(13)C composition determined. Although the extractives and the cellulose showed different absolute delta(13)C values, the patterns of change over time (as represented by the annual rings) were similar in most cases. Within an annual ring, carbon isotope ratios of extractives were correlated with the cellulose isotope ratio (R2 = 0.33 in sapwood, R2 = 0.34 in heartwood for aqueous acetone-soluble extractives; R2 = 0.41 in sapwood for hot-water-soluble extractives). These data suggest that some sapwood extractives are formed when the wood ring forms, and remain in place until they are converted to heartwood extractives many years later. Sapwood extractives appear to be important sources of materials for the biosynthesis of heartwood extractives in Douglas-fir.     

Intra-annual growth response of adult Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> [L.] KARST.) and European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) to an experimentally enhanced,free-air ozone regime

Several findings indicate an impact of ozone on stem diameter growth leaving the question unanswered, if and how the intra-annual growth pattern is changed. In this study the hypotheses are tested, that (1) ozone will alter the absolute growth and (2) alter and shift the period of growth activity within a year. Our data originates from the free air ozone fumigation experiment ‘Kranzberger Forst’ in a mixed stand of Norway spruce and common beech near Freising/Germany. Annual and intra-annual growth reactions of a sample of five adult beech and five spruce trees, exposed to double ambient ozone were examined and compared to the same number of untreated reference trees. Diameter increments were measured with plastic diameter girth bands and high-resolution, automatically logging micro-dendrometers, mounted at breast height (1.3 m). We used the increment data from the growth periods 2000 to 2005. The high-resolution micro-dendrometer data were examined by fitting a Weibull function to the standardized annual growth profiles to obtain curve parameters for statistical tests. We estimated the parameters ‘T’ which represents the point of time, when 63% of the annual diameter increment is performed and the parameter ‘m’, the Weibull module, which was used as an indicator for the span of time needed to complete the annual growth. The statistical significance of these curve parameters, together with the absolute diameter increment, was tested by use of mixed regression models. The analysis of the growth curve parameters revealed a significantly altered intra-annual growth pattern of both species induced by ozone. Spruce under ozone showed reduced absolute annual diameter increment and a preponed growth activity compared to untreated trees. Beech’s absolute diameter increment was not affected under ozone, but its growth activity was delayed. For both species, ozone fumigation did not alter the individual length of the annual growing season. These results are discussed with respect to drought, tree ring anatomy and tree allometry. The study shows that ozone is able to change growth behaviour of trees even if increment losses are not obvious. This article belongs to the special issue "Growth and defence of Norway spruce and European beech in pure and mixed stands".     

Seasonal nutrient dynamics in white pine and white spruce in response to environmental manipulation

Seasonal retranslocation in white pine (Pinus strobus L.) and white spruce (Picea glauca (Moench) Voss) was examined in response to silvicultural treatments (scarification, annual fertilization application, and annual control of competing vegetation with herbicide) that changed both environmental conditions and the growth rate of the trees. Four years after plantation establishment and initial treatment, nutrient accumulation in current-year needles of white pine and retranslocation from 1-year-old needles were increased following the vegetation control treatment, which increased resource availability (nutrients, water and light) and, hence, growth rate. Nutrient accumulation also increased in current-year white spruce needles following the same treatment, whereas retranslocation decreased in 1-year-old white spruce needles. Correlations of retranslocation (N, P and K) with growth rate (shoot biomass increment) showed a strong positive relationship for white pine and a negative relationship for white spruce. Retranslocation of K was correlated with foliar and soil K concentrations; the availability of this nutrient was also significantly reduced by vegetation control. A general theory for the control of nutrient retranslocation in conifers, which is not based exclusively on either sink strength or soil nutrient availability, is proposed. We conclude that retranslocation response is species specific and related to the potential phenotypic growth response to changing environmental conditions and to short-term imbalance in the supply versus the demand for nutrients.     

Some factors influencing susceptibility to discoloring fungi and water uptake of Scots pine (Pinus sylvestris), Norway spruce (Picea abies) and Oriental spruce (Picea orientalis)

Abstract

The heartwood and sapwood from Scots pine (PS), Norway spruce (PA), and Oriental spruce (PO) were tested for susceptibility to discoloring fungi and water uptake. In addition, annual ring width and density were measured. The methods used were Mycologg for testing growth of fungi and a modified version of EN 927-5 to investigate water uptake. For pine, the heartwood showed a lower water uptake and no discoloring fungi growing in the tests. The heartwood had a significantly higher density and smaller annual ring width than the sapwood. In PA the heartwood had significantly lower discoloration than sapwood. The total water uptake in g/m² was significantly higher in sapwood, but not the calculated moisture content. As for wood properties, the density was significantly higher in sapwood compared to heartwood, although there were no differences in annual ring width. Regarding PO, differences in water uptake could be seen between sapwood and heartwood although the densities were similar. These results show that susceptibility to discoloring fungi and water uptake is hard to correlate to a single inherent property when looking at different wood species.     

Forest decline: modeling the effect of climate in tree rings

Tree rings provide an historical record of forest growth that reflects changes with time in site factors including, competition, tree and stand age, fire and other disturbances, and climate. Statistical methods can be used to factor out climatic influences on radial growth to yield a climate response model that can indicate whether declines in forest productivity are related to the modeled climatic variables or to other influences such as atmospheric pollutants. A general method, based on ordinary least squares, is presented for creating climatic response models for forest decline studies. The crux of the method is model verification, whereby the time-stability of the model is tested before it is used to forecast tree-ring variations during a period of decline. Three studies are described that employ monthly mean temperatures to predict tree-ring indices in declining red spruce (Picea rubens Sarg.) stands in the Appalachian Mountains of North America. The results indicate that, since 1960, red spruce trees at most of the sites sampled have undergone a period of growth decline that is unrelated to changes in mean monthly temperature. However, an association between annual ring width and unusual departures from the mean summer and winter temperatures during both present and past periods of decline suggests that climatic effects are implicated to some degree in the current decline.     

Relationships among cold hardiness, root growth potential and bud dormancy in three conifers

Greenhouse-cultured, container-grown ponderosa pine (Pinus ponderosa var. scopulorum Engelm.), interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) and Engelmann spruce (Picea engelmannii (Parry) Engelm.) were cold acclimated and deacclimated in growth chambers over 19 weeks. Stem cold hardiness, total new root length at 14 days and days to bud break were measured weekly. Relationships among cold hardiness, root growth potential (RGP) and bud dormancy suggest that cold hardiness, which can be measured quickly, could provide a useful basis for estimating the two other parameters. During cold acclimation, there was a lag period in which stem cold hardiness remained at -15 degrees C and RGP was at a minimum, in all three species. Douglas-fir and Engelmann spruce buds remained fully dormant during this lag period. Ponderosa pine buds had no chilling requirement for the loss of dormancy, and reached quiescence during the lag period. Immediately following the lag period, as stem cold hardiness progressed to -22 degrees C, RGP increased to a high plateau in all three species, and Douglas-fir and Engelmann spruce buds approached quiescence. Cold deacclimation and bud development began immediately on exposure to warm, long days, but RGP remained high until stem cold hardiness returned to approximately -15 degrees C. At bud break, cold hardiness and RGP were at the minimum.     

Tall planting stock for enhanced growth and domination of brush in the Douglas-fir region

Two long-term experiments followed development of planted stock of various sizes, origins, and species. In one experiment, multiyear comparisons of container, 2-0 bareroot, and 3-year-old Douglas-fir transplants showed strong positive relationship between initial height and long-term (10–14 years) growth under a range of site conditions with high probability of brush development. In the other experiment, Douglas-fir, western hemlock, and Sitka spruce were planted in salmonberry disturbed by logging 0 and 4 years previously. Half the seedlings were released with glyphosate 6 months after planting. Hemlock and Douglas-fir bare-roots all grew well if planted in a fresh burn, despite rapid regrowth of salmonberry, but virtually all seedlings less than 60 cm tall except Sitka spruce were killed by 4-year-old salmonberry if not released. Release improved growth of seedlings in the fresh burn by 6%, gaining an average of about 0.6 year toward reaching a height of 6 m. Release improved growth of survivors in 4-year-old salmonberry by 51% in height, 72% in diameter, and 325% in volume at age 12. Sitka spruce grew well until damaged by insects. Hemlock growth was equal to or greater than that of Douglas-fir of comparable initial height. In all comparisons, the probability of being overtopped by brush decreased with increasing initial stock height, and the effect of suppression on growth was also inversely related to initial height. Field planting operations may require special logistic measures for the largest stock types.This is Paper 2706 of the Forest Research Laboratory, Oregon State University.     

What is hot in tree rings? The wood density of surviving Douglas-firs to the 2003 drought and heat wave

In a global warming scenario, drought and heat waves like the one that occurred in 2003 in Europe are expected to become more intense and frequent. This extreme climate event strongly affected the hydraulic balance in many forest tree species including Douglas-fir, with symptoms ranging from partial crown necrosis to death. We studied a French Douglas-fir provenance trial, strongly affected by the 2003 drought and heat wave. Using wood X-ray microdensity profiles as a record of cambium response to environmental variations, we compared mean stem density and growth between dead and alive neighbouring trees and several microdensity characters measured over 17 growth rings previous to 2003. Special attention has been given to the sampling layout in order to minimize the micro-environmental effects. At tree level, surviving trees have a significantly higher mean stem density. At tree ring level, they have a significantly higher mean ring density, maximum ring density, latewood density and, to a lesser extent, latewood proportion. No significant difference was found for diameter growth. These results could have direct implications in the way to study and predict possible acclimation and adaptation of forest trees to climate change process. Wood could be used for the selection of genotypes with a desirable plasticity and resistance to drought induced-cavitation.     

Classification of Picea abies Pulpwood According to Wood and Stand Properties

Different classification systems for Norway spruce (Picea abies (L.) Karst.) pulpwood were compared. The classification systems were applied on truckloads or single logs in southern Sweden. Truckload classification according to mean annual growth ring width gave better separation of the wood properties basic density, juvenile wood and dry matter content, than classification according to harvest type (first thinning, later thinning or final felling). The assortments did not have significantly different wood brightness. Sorting at log level according to diameter, mean annual growth ring width or number of annual growth rings, which could be done at harvesting, did not drastically improve differentiation of the mean values of the wood properties or reduce variance compared to truckload classification. The variation in wood properties within assortments remained large owing to the large variation in wood properties between and within logs. Substantial reduction in dry matter variation could be achieved by truckload classification during the summer.     

Blight of conifer seedlings caused by Colletotrichum gloeosporioides

In a growth chamber experiment, 70-day-old seedlings of 10 conifer species were inoculated with Colletotrichum gloeosporioides conidia to determine the host range of the fungus. Based on the percentage of seedlings affected and the disease severity on individual seedlings, the order of most to least susceptibility was: western hemlock (WH), mountain hemlock, western larch, Sitka spruce, Engelmann spruce, Douglas-fir (coastal form, then interior form), white spruce and ponderosa pine; lodgepole pine and western red cedar remained unaffected. Inoculation of WH needles showed that within 24 h C. gloeosporioides conidia germinate and appressoria (penetration structures) form. A growth chamber study demonstrated that the pathogen can infect WH at needle wetness periods as short as 15 min; number of needles affected was higher at 0.5 h, but did not increase further even when wetness was extended up to 8 h. The results are discussed in relation to blight management of greenhouse-grown conifer seedlings.Portion of a Bachelor of Science (Honors) thesis, Department of Biology, University of Victoria, Victoria, British Columbia.     

Indented rings (hazel growth) of Norway spruce reduce anisotropy of mechanical properties

This study presents three-point bending test results of Norway spruce clear wood specimens loaded on the radial-longitudinal plane in two different load cases. The tested samples were graded as resonance wood for instrument making and were characterised by narrow annual rings and relatively low density. The modulus of elasticity (MOE) and the corresponding modulus of rupture (MOR) are illustrated separately for the samples with straight grain and the group showing the specific growth pattern of indented rings (‘hazel growth’). With the longitudinal wood anatomical direction parallel to span width, the fibre deviation caused by the indents reduces MOE and MOR values, whereas a ‘reinforcing’ effect of the indents could be observed for the load case with span width parallel to the radial direction. Both aspects lead to a reduction in anisotropy for hazel-growth Norway spruce (anisotropy MOE: indented rings 11.6, straight grain 14.7, anisotropy MOR: indented rings 6.9, straight grain 8.9), which partly explains the exceptional position of this growth pattern for the construction of high-class musical instruments with outstanding mechanical and acoustical performance.     

Responses of Picea, Pinus and Pseudotsuga roots to heterogeneous nutrient distribution in soil

The spatial distribution of plant-available mineral nutrients in forest soils is often highly heterogeneous. To test the hypothesis that local nutrient enrichment of soil leads to increased root proliferation in the nutrient-rich soil zone, we studied the effects of nutrient enrichment on the growth and nutrient concentrations of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) roots. Three-year-old seedlings were grown for 9 months in split-root containers filled with nutrient-poor forest mineral soil, with one side supplemented with additional mineral nutrients. Root dry weight and root length in Scots pine and Norway spruce were increased in the nutrient-supplemented soil compared with the nonsupplemented side, whereas root growth in Douglas-fir was unaffected by nutrient enrichment. Of the three species examined, Norway spruce exhibited the highest root and shoot growth and the highest nutrient demand. Specific root length (m g(-1)) and the number of root tips per unit root length were not affected by local nutrient addition in any of the species. Despite increased root growth in Norway spruce and Scots pine in nutrient-supplemented soil, their root systems contained similar nutrient concentrations on both sides of the split-root container. Thus, coniferous trees may respond to local nutrient supply by increased root proliferation, but the response varies depending on the species, and may only occur when trees are nutrient deficient. As a response to local nutrient enrichment, increases in root dry matter or root length may be better indicators of pre-existing nutrient deficiencies in conifers than increases in root nutrient concentrations.     

Optimal timing of early genetic selection for sawn timber traits in <Emphasis Type="Italic">Picea abies</Emphasis>

In breeding Norway spruce, selection for improved growth and survival is performed at age 10–15 years in order to optimize genetic gain per year. We investigated whether a selection based on wood traits such as density and grain angle, measured under bark in the field at the same age would be informative enough with respect to structural quality traits of sawn boards. To achieve this objective, a sawing study was conducted on the butt logs of 401 trees from a 34-year-old Norway spruce progeny trial situated in southern Sweden. Stem discs were excised from the top of the logs and radial profile data of grain angle, and wood density was recorded for specific annual rings. The sawn and dried boards were assessed for structural traits such as twist, board density, bending stiffness (static modulus of elasticity, sMoE) and bending strength (modulus of rupture, MoR). Additive genetic correlations (r_a) between single annual ring density measurements and board density, sMoE and MoR were consistently strong (r_a>?0.7) for annual rings 5–13. Genetic correlations of similar magnitude between grain angle and board twist were estimated for all investigated annual rings (from 2 to around 26 under bark). Consequently, it was found that indirect selection for wood density and grain angle at the tree age 10–16 years would result in more genetic gain per year than selection at later ages. This makes it feasible to perform simultaneous selection of progeny in the field for both growth and wood traits at similar ages.     

Coarse root elongation rate estimates for interior Douglas-fir

Accurate estimates of root growth rates are important for root system modeling, and the spread of root systems may be an important determinant of belowground site occupancy. Estimating root system growth rates is complicated because missing, discontinuous, and false annual growth rings make root cross sections difficult to age. These irregularities can occur even in roots of dominant conifers with rare or absent stem growth ring abnormalities. Incomplete rings were noted in the root growth rings of nine co-dominant interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) trees. Coarse root (> 1.0-cm diameter) elongation rates were estimated by fitting a geometric mean regression line to ring count and lateral distance data. In all nine roots examined, the geometric mean regression slope was well within the range of the 95% confidence interval for the ordinary least squares regression of lateral distance versus age, suggesting that measurement error may have been negligible. Coarse root elongation rates (which ranged from 2.8 to 15.3 cm year(-1) and averaged 7.4 cm year(-1)) in the interior Douglas-fir trees studied were much lower than those reported by others. This discrepancy may be a result of limited soil water availability, soil heterogeneity (both soil water content and soil texture were highly variable across short distances) and fragmentation of belowground growing space.     

Simultaneous X-ray diffraction and X-ray fluorescence microanalysis on secondary xylem of Norway spruce

Secondary xylem of Norway spruce was studied by X-ray microanalysis. Average dimensions of cellulose crystallites, fraction of oriented cellulose, mean microfibril angle, and nutrient element concentrations of K, Ca, Mn, and Zn were simultaneously determined using microfocused synchrotron radiation and a combination of X-ray diffraction and X-ray fluorescence spectroscopy techniques. The variation of these quantities in the microscopic size scale was noticeable, and similar between samples taken from the same annual ring. The mean microfibril angle and the nutrient concentrations of Ca, Mn, and Zn showed a correlation. The mean values of the structural parameters and their variation as a function of the annual ring were similar as reported in previous studies on Norway spruce.