Regeneration patterns of Eastern white pine (Pinus strobus L.) in hardwood-dominated forests in Virginia, USA

Eastern white pine (Pinus strobus L.) is a moderately shade-tolerant species that co-occurs with hardwood tree species in many forests of the eastern United States, as well as in pure stands. The species is valued for its timber, as well as for wildlife and recreation. Regeneration of this species is somewhat unpredictable and often occurs in patches of similarly-aged cohorts. We described the regeneration patterns of this species and examined their relation to environmental variables within hardwood forests of southwestern Virginia, USA. An average of 5.3 white pine patches per ha were observed in this study. The majority of patches consisted of saplings (85%), with 9% of patches in pole size classes, and 6% in seedling size classes. The average density of patches was 43.5 stems with an average age of 20 years. The size of patches averaged 80.6 m². The total density of seedlings and the number of regeneration patches of all sizes of regeneration (seedlings, saplings, and poles) in plots was related to the surrounding density of large white pine trees (potential seed trees). The density of seedlings or patches was not significantly related to current vegetation cover or soil surface cover variables, but more than half of regeneration patches were located in or adjacent to old canopy gaps, most of which were old logging gaps. While seedling regeneration may occur within the understory of these forests near seed trees, advancement to the sapling and pole stage appears to be associated with canopy gap formation.     

Comparative mast seed production in unmanaged and shelterwood white pine (Pinus strobus L.) stands in central Ontario

The effect of the seed cut of the uniform shelterwood silvicultural system on white pine seed production, seed characteristics, and seed viability during 2 mast seeding events was examined in operationally harvested second growth, white pine-dominated forest stands in central Ontario. Seed traps placed along transects in unmanaged and shelterwood stands in each of 3 blocks were used to monitor seed production in 2000 and 2006 (4 and 10 years after harvesting). During these 2 mast seed years between 386,000 and 2,730,600 seed ha^?1 were produced among study stands. Total seed production expressed on a per hectare and unit pine basal area basis did not differ by harvest treatment or among blocks in either year. Variability in seed production among stands was primarily due to differences in stand structure, with seed production positively related to white pine basal area. Seed characteristics were largely similar between harvested and unmanaged stands and between seed years. Seed viability was relatively high in both years, with seed from shelterwoods germinating slightly slower than those from unmanaged stands. Seed quality, as estimated by laboratory germination performance, was higher in 2006 than 2000, likely due to improved seed development and maturation in the warmer, wetter growing season of 2006. Our results suggest that the seed cut of the uniform shelterwood system applied to second growth white pine stands is unlikely to adversely affect white pine seed production, seed quality, or potential for natural regeneration during mast seeding events.     

Effects of soil organic matter,moisture, shading and ash on white pine (Pinus strobus L.) seedling emergence

The effects of soil organic matter, soil moisture, shading and ash on white pine (Pinus strobus L.) emergence were investigated using soil monoliths in greenhouse experiments. White pine seedling emergence increased with soil organic matter removal, and levelled with the elimination of the litter and fermentation layers, where seedling emergence did not significantly increase further. Increased shade and soil moisture levels also improved seedling emergence whereas ash had no or little impact on white pine seedling emergence. The silvicultural implications of these findings are discussed with regard to white pine management.     

Natural Variation in Seed Characteristics of Eastern White Pine (Pinus strobus L.)

Natural variation in seed characteristics of eastern white pine (Pinus strobus L.) trees and its relationship with the size, form, and crown class of parent trees was examined in second-growth, white pine mixedwood stands in Ontario. Cones were collected prior to and following partial harvesting of two stands during 2 years of above-average seed production. Fully enlarged seed extracted from mature cones was X-rayed to determine seed yield per cone, the percentage of sound, partially filled, and empty seed, and the sound seed mass for each tree. Variation in seed yield per cone, the proportion of sound, partially filled, and empty seed, and seed mass was much greater among individuals than between stands or seed years. The strength and significance of the relationship of seed mass with sound seed yield differed with stand and seed year. In uncut stands, seed mass was higher for trees with dominant crown class, larger live crown ratios, and smaller height to diameter ratios, suggesting tree vigour and microsite quality influence seed mass. Proportion of partially filled seeds was significantly reduced in both stands in the second year likely due to increased growing degree days during female gametophyte development. The implications of these results to natural and artificial regeneration of white pine are discussed.     

Microstructural investigation of UV-laser irradiated pine (Pinus silvestris L.)

Summary Different surfaces of pre-sanded pine (Pinus silvestris L.), namely radial surfaces, tangential surfaces and cross sections, have been subjected to microstructure examination. UV-laser irradiation with a wavelength of 248 nm and an energy of 350 mJ is used to enhance the microstructure exposure. It has been shown that UV-laser irradiation is a useful and practical method for revealing microstructural details. This is more pronounced on the cross section. Sanding with emery paper finer than 800 mesh can reveal microstructure on the radial surface equivalent to that revealed by UV-laser irradiation. On the radial and tangential surfaces, more earlywood is extracted after UV-laser irradiation, resulting in an uneven surface. Some features such as loose fibres and a rack-gearing-like structure on the radial and tangential surfaces, as well as a loss of radially oriented cell walls on the cross section are observed as a result of the UV-laser treatment. The rack-gearing structure is composed mainly of a sawtooth-like pattern. The length of the sawtooth is in most cases approximately 9 and 19 m.This work was carried out in the Department of Wood Technology and Processing, Royal Institute of Technology (KTH). Financial support from KTH is gratefully acknowledged.     

Histopathology of primary needles and mortality associated with white pine blister rust in resistant and susceptible Pinus strobus

White pine blister rust caused by Cronartium ribicola is a damaging non‐native disease of five‐needled pines in North America. Efforts to control the disease and mitigate damage to date have been only somewhat effective. Recent efforts to improve the health of eastern white pine and reestablish the tree as a dominant species in the North Central United States have focused on identification and propagation of disease‐free eastern white pine (Pinus strobus) growing in areas with a high incidence of blister rust. Many of these selections have been shown to resist infection following artificial inoculation with C. ribicola. In this study, 13 eastern white pine families derived from controlled pollination of selections previously determined to possess putative resistance as well as susceptible selections were inoculated with C. ribicola. Mortality data from inoculation studies show superior survivability in three families with over 60% of seedlings able to survive the 52 week post‐inoculation monitoring period compared to 0–10% survival of the most susceptible families. Primary needles were collected for histological analysis from all inoculated families 4 weeks after inoculation and from selected families 6.5 weeks and 38 weeks after inoculation. Histological observations of infection sites show distinct resistance reactions in the families more likely to survive infection based on mortality data. Analysis of the reactions in susceptible families revealed extensive hyphal colonization of the vascular bundle and adjacent mesophyll cells that appear uninhibited by tree responses. In resistant families, collapsed cells adjacent to infection sites, heavy deposition of phenolic compounds and abnormal cell growth were documented more frequently and appear to play an integral role in the ability of these eastern white pine families to impede growth of C. ribicola in primary needle tissue.     

Crown architecture and stemwood production in Norway spruce (Picea abies (L.) Karst.)

The morphological data for Norway spruce (Picea abies (L.) Karst.) trees published by Burger (1953) were reanalyzed to examine the relationships between crown shape, needle and branch mass density of crown volume, needle efficiency in stemwood production, harvest index and stemwood production efficiency per unit of crown projected area. Production of stemwood per unit of crown projected area was higher in narrow-crowned trees than in broad-crowned trees because the narrow-crowned trees had (i) smaller horizontal space requirements for equal crown volumes, (ii) higher leaf area index due to both the geometrical crown shape and higher needle density per unit of crown volume, and (iii) higher harvest index indicating high allocation of dry matter production to stem.     

Mevalonate kinase activity during different stages of plant regeneration from nodular callus cultures in white pine (Pinus strobus)

Mevalonate kinase (MK) catalyzes a step in the isoprenoid biosynthetic pathway, which leads to a huge number of compounds that play important roles in plant growth and development. Here, we report on changes in MK activity in white pine (Pinus strobus L.) during plant regeneration by adventitious shoot organogenesis from cotyledons of mature embryos, including nodular callus induction, shoot formation and rooting. Nodular calli were induced from Pinus strobus (PS) embryos by culture in nodular callus induction medium in a 0-, 8- or 16-h photoperiod. Mevalonate kinase activity peaked in nodular calli after three weeks of culture on nodular callus induction medium in a 16-h photoperiod, whereas frequency of nodular callus formation peaked after 4 weeks of culture on nodular callus induction medium in darkness. During adventitious shoot formation, MK activity peaked in shoots derived from dark-grown nodular calli after 3 weeks on bud formation medium, and frequency of shoot formation was highest in dark-grown nodular calli cultured on bud formation medium for 4 weeks. During rooting, MK activity peaked 2 weeks after transfer of adventitious shoots to rooting medium and rooting frequency was highest in adventitious shoots after 3 weeks on rooting medium. Although during nodular callus induction in darkness MK activity was inversely related to frequency of nodular callus formation, MK activity was highly correlated with frequency of shoot formation and with rooting frequency. The observed increase in MK activity preceding rooting suggests that MK could serve as a marker for rooting of white pine shoots in vitro.     

Presence of longitudinal cracks in planks from storm-felled pine (Pinus sylvestris L.) and spruce (Picea abies (L.) Karst.)

Abstract

After the severe storm Gudrun in southern Sweden in 2005, a quantitative study was done in order to investigate the presence of lengthwise crack on planks taken from storm-felled trees in southern Sweden, compared to planks from standing trees not subjected to this storm (central Sweden). The main yield from each log was examined. In total, 1087 pine (Pinus sylvestris) planks and 3626 spruce (Picea abies) planks from the storm-struck area were investigated and compared to 1953 spruce and 2000 pine planks from trees outside the storm-struck area. The examination of cracks was done visually on dried planks. For pine, 51.7% of the planks from storm-felled trees had a total length longer than 0.5 m, compared to 7.3% for the reference material. As for spruce, 11.0% of the planks from storm-felled trees had a total crack length of more than 0.5 m, compared to the reference material where 2.2% had cracks longer than 0.5 m. The results show that the storm-felled trees had more longitudinal cracks than the reference material and that pine was more likely to develop storm-related cracks than spruce.     

Crookedness of pedunculate oak (Quercus robur L.) growing under a canopy of Scots pine (Pinus sylvestris L.)

Natural regeneration of pedunculate oak growing under a canopy of Scots may be used for conversion into a mixed or predominantly broadleaved forest. From an economic perspective it is desirable that the oaks have straight stems. In this paper we present a case study for analysing oak stem curvature and its causes. The study was conducted in a mature pine stand with understorey oaks of age ca. 60 years in Poland. The oaks were classified into two main groups as either straight or crooked. The following variables were measured or assessed for the oaks: (1) stem curvature (crookedness), total tree height, diameter at breast height (DBH) and some tree crown characteristics, (2) the growth of DBH and height in each year of the life of each tree and (3) the competitive pressure of surrounding trees. Durable stem curvatures were formed at the culmination of the height increment. The first 20 years of the life of the oak were crucial in terms of stem quality. The annual height increment was larger in the years when the curvature was formed than in adjacent years. The factors that caused crooked stems were the same for straight and crooked oaks, but the magnitude of the response was different. When planning the conversion of Scots pine into oak, full overhead light should be provided as early as possible, but not later than 20 years since the establishment of the regeneration.     

Development and transferability of two multiplexes nSSR in Scots pine (Pinus sylvestris L.)

Single sequence repeat(SSR) multiplexing is a semi high-throughput PCR methodology for the analysis of multiple SSRs.We developed two SSR multiplexes selected from SSR loci previously reported in the pine literature and tested the transferability of both SSR multiplexes in nine other pine species.We tested 234 nuclear SSR loci(n SSRs) previously described in the pine literature and selected ten n SSRs following the simple criteria of interpretability and reproducibility.Selected nuclear loci were divided into two n SSRs multiplex sets and their amplification was optimized for three different multiplex PCR methods based on:(a) a custom PCR protocol,(b) a custom protocol with hotstart taq polymerase,and(c) a commercially available kit for SSR multiplexing.To validate their performance,the level of genetic diversity was assessed in three Scots pine natural populations(Hungary,northern Sweden and southern Sweden).In addition,we also tested the transferability of these multiplexes in nine other pine species.We have developed two n SSRs multiplexes of five loci each that will contribute to reduce the costs of n SSR scoring,while increasing the capacity of n SSR loci analysis.Amplification was successful in all three populations(94 % success) and the level of polymorphism(7.1 alleles/marker) was similar to that previously reported for other Scots pine natural populations.Transferability of both multiplexes was successful for those pine species closely related to Scots pine.     

Estimating sapling vitality for Scots pine (Pinus sylvestris L.) in Russian Karelia

A new method is proposed for estimating vitality or growth potential for saplings of Scots pine (Pinus sylvestris L.), based on height, diameter and height increment. A two-stage process was used to establish the vitality index. The logarithms of height, diameter and height increment were regressed against age, to adjust for the wide range of ages present in our data (c. 10,000 saplings with ages spanning 4–50 years). Then principal component analysis was used to obtain coefficients, which were, in turn, standardized on each axis to provide a vitality index scaled in standard deviations. This standardized scale allows the rank of an individual in the population to be assessed, and draws attention to possible outliers. The use of age-adjusted residuals ensured that the estimator was independent of age, and stable over a wide age range. The first principal component indicates if a sapling is relatively tall (weight = 0.5), thick (w = 0.5) or fast-growing (w = 0.7) for its age. Most of the information is contained in the first principal component, but the second component, which explains about 10% of the variance, appears to offer some utility as an indicator of ‘acceleration’ due to changing conditions. The resulting measures of vitality have been useful for research and management in the dry lichen-moss pine forest in Russian Karelia, but are specific to this species, locality and ecotype. Further research and site-specific data are necessary to adapt the system to other situations.     

Fine root respiration in mature eastern white pine (Pinus strobus) in situ: the importance of CO(2) in controlled environments

We measured seasonal fine root respiration rate in situ while controlling chamber temperature and [CO(2)]. Atmospheric [CO(2)] ([CO(2)](a)) and measured soil [CO(2)] ([CO(2)](s)) were alternately delivered to a cuvette containing intact fine roots of eastern white pine (Pinus strobus L.). Respiration rates were consistently higher in [CO(2)](a) than in [CO(2)](s) and were almost three times higher during midsummer. Respiration rates were immediately reversed after returning to the alternate [CO(2)] (i.e., [CO(2)](a) --> [CO(2)](s) --> [CO(2)](a), and vice versa) suggesting a direct effect of elevated [CO(2)] on apparent respiration. Soil-[CO(2)]-based respiration rates decreased with increasing [CO(2)] on a dry mass and tissue [N] basis. We conclude that estimates of soil CO(2) flux and soil carbon budgets may be improved by more completely accounting for the rhizosphere microclimate (i.e., soil temperature and [CO(2)](s)) during measurement of fine root respiration.     

Effect of magnetic treatment on seed germination of loblolly pine (Pinus taeda L.)

Based on preliminary experiments, the speed of germination (SG) was in general increased for Pinus taeda L. seeds treated with a static magnetic field (B = 150 mT) for 10, 30 and 60 min. Negative impact was obtained for seeds treated with SMF for 24 h and 48 h. Mean germination time (MGT), the SG, and time required to obtain 10%, 25%, 50%, 75%, and 90% of seeds to germinate (T₁₀–T₉₀) were calculated. Results showed a reduction of the MGT and T₁₀–T₉₀ for seeds treated with SMF for 10, 30, and 60 min; therefore, the germination speed was increased. Among various magnetic treatments, loblolly pine seeds treated with S3 (SMF60 min) yielded the peak performance.     

Wood quality assessment of Pinus radiata (radiata pine) saplings by dynamic mechanical analysis

Wood Science and Technology - The use of dynamic mechanical analysis was explored as a possible method of screening for wood quality in breeding programmes. Viscoelastic properties along the grain...     

Righting response of artificially inclined maritime pine (Pinus pinaster) saplings to wind loading

To determine if trees respond to dynamic and static loading in the same manner, 2-year-old maritime pine (Pinus pinaster Ait.) trees were subjected to different types of mechanical loading in the field. One block of trees (the control) were kept in pots and planted in the field at an angle of 0 or 45 degrees to the vertical. A similar block of leaning potted trees was planted nearby and subjected to frequent, unilateral wind loading for a period of 1 s every 2 min. Half the leaning trees were oriented toward the direction of wind loading and half were oriented along the axis of wind loading. The stem profile was measured three times during the growing season to quantify the rate of stem straightening. Compression wood formation and stem shape were measured in all plants. No differences in mean height or diameter were observed between blocks and all leaning trees straightened, but not at the same rate. Although no difference in the rate of apical straightening occurred between control and wind-treated trees, the righting response of the basal part of the stem of leaning trees subjected to wind was four times greater than that of leaning trees without wind. No differences in the righting response were observed between leaning trees growing toward and trees growing away from the source of wind. No significant differences in compression wood formation were found between control trees and wind-treated trees, indicating that other factors must determine the reorientation rate of leaning trees. Results are discussed with reference to the quality of compression wood in conifers and the mechanotransductive pathway in plants.     

Dynamic response of stomata to changing irradiance in loblolly pine (Pinus taeda L.)

Dynamic changes in stomatal conductance and the rate of photosynthesis were measured as periods of shading (decrease in irradiance from 800 to 200 micro mol m(-2) s(-1)) between 5 and 60 min were imposed on needles of Pinus taeda L. trees under laboratory conditions. Shading induced a 39% decrease in stomatal conductance but the rate of change was slow. Average time constants (+/- standard error) were shorter for the decrease in stomatal conductance when shading was imposed for 30 min (14.8 +/- 1.3 min) than for the increase in stomatal conductance when shading was removed (25.5 +/- 3.4 min). The time constants for increasing stomatal conductance when shading was removed were linearly related to the length of the previous dark period. The rate of photosynthesis fell immediately by 58% when shading was imposed and increased more rapidly than the change in stomatal conductance when shading was removed. The increase in photosynthesis during the induction phase after shading was removed was limited by both stomatal and biochemical effects. The long time constants for stomatal response contributed to the poor correlations between stomatal conductance and instantaneous measurements of irradiance from field data. However, the slow response of stomatal conductance to changes in irradiance had little effect on total daily transpiration, carbon gain and water-use efficiency.     

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.