A comparison of red spruce and balsam fir shoot structures

I compared the shoot structures of high-elevation red spruce (Picea rubens Sarg.) and balsam fir (Abies balsamea (L.) Mill.). Needle widths, thicknesses and perimeters were measured to estimate total leaf areas from measured projected leaf areas. Measured needle perimeter/needle width ratios differed significantly from estimated ratios that assumed needles were either rhomboidal or elliptical in cross section. The vertical and horizontal silhouette shoot area to total leaf area ratios (STAR(v) and STAR(h)) of the two species were negatively correlated with needle packing and canopy height. Red spruce had higher values of STAR(v) than balsam fir at each canopy height, but STAR(v) declined with canopy height at a similar rate in the two species. The STAR(h) values of the two species did not differ significantly at a given canopy height. Needle packing increased with canopy height at the same rate in the two species. Needle weight increased in red spruce and decreased in balsam fir with increased needle packing, but showed no significant dependence on canopy height. Red spruce had higher values of STAR(h) than balsam fir at low values of needle packing, but STAR(h) values converged at high values of needle packing. The generally comparable values of STAR, along with similar needle diameters, may imply that red spruce and balsam fir have similar collection efficiencies of wet and dry particles. Measurements of STAR may be used to estimate leaf area indices (LAI) more accurately when using indirect techniques.     

Diffusion of dissolved oxygen in liquid-saturated Douglas fir sapwood

Summary A technique was developed for the determination of the effective dissolved oxygen diffusivity in liquid-saturated softwood in a diffusion cell under ambient pressure. From the measurements in the temperature range 2° to 50°C, the activation energy of diffusion was found to be 4.6 kcal/mole. The diffusivity in summerwood is about one half of that in springwood. The diffusivity of dissolved oxygen through composite springwood and summerwood layers was based upon samples of two Douglas fir sapwood blocks saturated with water. In the radial and tangential directions, diffusivity was 1.4 to 2.3x10^-6cm²/s, which is about 6 to 10% that of dissolved oxygen diffusivity in water. The diffusivity in the longitudinal direction is about 5.5 times that in the other two directions.The diffusivity of dissolved oxygen through liquor-saturated wood and the effect of delignification on oxygen diffusion were also determined.The authors are grateful to Weyerhaeuser Company and Air Products and Chemical Inc., for the financial support of this research.     

An effect of gravity on bud-burst in balsam fir

Bud-burst on first order lateral branches of Abies bafsamea L. (balsam fir) was delayed when the branches were rotated 180 degrees about their long axis. This was not a consequence of injury caused by the treatment because buds rotated 180 degrees on inverted plants flushed at the same time as the controls, whereas flushing of all other buds was delayed. Buds thus appear to be more vigorous when maintained in the same orientation to gravity in which they are formed and the site of gravitational stimulus perception appears to be the bud itself. Except on the leading shoot, leaves from inverted buds turned so that their adaxial surface faced upward, unless there was intense illumination from below. However, both anisophylly and positioning of leaves on lateral shoots were apparently predetermined because the shorter, more forward pointing leaves appeared below the longer distichous leaves on shoots from inverted buds. Shoots with normally oriented leaves appeared the next season.     

Response of balsam fir and hemlock roots to injuries

Drill wounds in balsam fir and hemlock roots activated the nonspecific resistance mechanisms of compartmentalization in wood and necrophylactic periderm in bark. Tangential bands of resin ducts localized around the wounds constituted the barrier zones in the secondary xylem of conifer roots. Barrier zones were more extensive in roots which showed symptoms characteristic of invasion by fungi and bacteria after wounding. This observation supports an expanded definition of barrier zones; barrier zones may form not only in response to mechanical wounds but also in response to xylem injury caused by pathogens. Multiple bands of resin ducts were common in young xylem when bark lesions developed around wounds. Necrophylactic periderms isolared dead bark from living bark. Development of phellem cells with dark contents and thick suberized walls, typical of exophylactic periderm, followed initiation of necrophylactic periderm. The wound responses were similar in both balsam fir and hemlock.     

Estimating cumulative defoliation of balsam fir from hemlock looper and balsam fir sawfly using aerial defoliation survey in western Newfoundland,Canada

Both hemlock looper (Lambdina fiscellaria fiscellaria (Guen.)) and balsam fir sawfly (Neodiprion abietis (Harris)) undergo periodic outbreaks in eastern Canada and cause significant growth and mortality losses to forests. Tree growth and mortality are closely related to cumulative defoliation estimates, which integrate annual defoliation over multiple years. Our objective was to determine a method to estimate cumulative defoliation of balsam fir (Abies balsamea [L.] Mill) due to these insects in western Newfoundland, using aerial defoliation survey data, as an essential input to modeling impacts for Decision Support Systems. Interpretation of aerial defoliation survey data for hemlock looper and balsam fir sawfly is problematic because both insects feed upon multiple age classes of foliage. Current-year (2008) aerial defoliation survey data were compared with ground estimates of defoliation by age class from 45 plots (450 trees and 395 mid-crown branch samples), representing a range of defoliation severity classes for each insect. Cumulative defoliation was calculated using defoliation per foliage age class, weighted by relative foliage mass for a given age of foliage. Three significantly different severity classes were defined based on cumulative defoliation values derived from aerial defoliation survey: (i) 1-year moderate (30–70%) defoliation, (ii) 1-year severe (71–100%) defoliation with calculated cumulative defoliation values of 19 and 39%, respectively, for balsam fir sawfly, 21 and 34% respectively for hemlock looper; and (iii) 2–3 years of moderate–severe defoliation, with cumulative defoliation ranging between 59 and 64% for balsam fir sawfly and 49% for hemlock looper. Defoliation severity from aerial defoliation survey alone hence can be misleading if defoliation measurements are not converted to cumulative defoliation values.     

Broom rusts of balsam fir and black spruce in Newfoundland

The broom rusts of balsam fir and black spruce occur sporadically throughout the island of Newfoundland, but they are not a serious threat to the forests. The incidence and intensity of the rusts vary, but no tree mortality can be attributed to the diseases. The average number of brooms per tree was higher in black spruce than in balsam fir. Also, more brooms occurred on brandies than on tree trunks. Height and diameter growth was less in infected trees than in uninfected trees of both the species.     

Separation of eastern spruce and balsam fir by chemical methods

Summary Increased utilization of eastern spruce and balsam fir has led to a need for a quick method of separating these woods in a mill situation. One such method might be the use of a chemical indicator. A test of various classes of chemical agents applied at different seasons of the year showed that a pH indicator might be suitable for achieving a separation. The most suitable indicator was tested on samples from different geographic locations and at three highproduction stud mills. Additional tests were conducted to explore such variables as moisture content and surface condition of the wood, type of solvent, concentration and temperature. The most suitable indicator found was bromophenol blue at a concentration of 0.10 percent in 95 percent ethanol. When applied to green wood which had been allowed to dry for a few minutes to a few hours, this indicator produced various shades of orange, yellow, green or blue with spruce and a dark blue or blue-violet with fir. With an understanding of the variables that affect the reaction, it is felt that bromophenol blue can be used for the separation of eastern spruce and balsam fir on a commercial basis.The authors would like to thank fellow researchers in New York, Vermont, New Hampshire and New Brunswick, Canada, for providing samples for testing. Also, assistance from the following stud mills in Maine is greatly appreciated: Diamond International Corporation (Passadumkeag), Georgia-Pacific Corporation (Woodland) and St. Regis Paper Company (Costigan). This research was financed under McIntire-Stennis Research Project 5-5-39602.     

Estimating stem maintenance respiration rates of dissimilar balsam fir stands

Stem maintenance respiration rates were measured in five contrasting balsam fir (Abies balsamea (L.) Mill.) stands. At 15 degrees C, average respiration rates for individual stands ranged from 120 to 235 micro mol m(-3) s(-1) when expressed per unit of sapwood volume, from 0.80 to 1.80 micro mol m(-2) s(-1) when expressed per unit of stem surface area, and from 0.50 to 1.00 micro mol g(-1) s(-1) when expressed per unit of nitrogen in the living stem biomass, but differences among stands were not statistically significant. Coefficients of variation ranged from 50 to 100% within stands and were similar for all bases used to express respiration rates. Coefficients of determination for regressions between chamber flux and chamber values of sapwood volume, stem surface area and nitrogen content varied between stands and no one base was consistently higher than the other bases. We conclude that the bases for expressing stem respiration are equally useful. Respiration rates were more closely correlated to stem temperature observed approximately 2 h earlier than to current stem temperature. Among stands, annual stem maintenance respiration per hectare varied from 0.1 to 0.4 Mmol ha(-1) year(-1), primarily because of large differences in sapwood volumes per hectare. Annual stem maintenance respiration per unit of leaf area ranged from 3 to 6 mol m(-2) year(-1), increasing as sapwood volume per hectare increased.     

Springtime resumption of photosynthesis in balsam fir (Abies balsamea)

Photosynthesis in balsam fir (Abies balsamea (L.) Mill.) was measured in the field at two locations in New Brunswick, Canada from late winter to late spring in 2004 and 2005. No photosynthesis was detectable while the soil remained below 0 degrees C throughout the rooting zone. In both years, photosynthesis began once soil temperature rose to 0 degrees C. In potted seedlings in growth chambers, there was no photosynthesis at an air temperature of 10 degrees C if the pots were frozen. These findings suggest that, once air temperatures permit photosynthesis, it is the availability of unfrozen soil water that triggers the onset of photosynthesis. In the field, full recovery of photosynthetic capacity following the onset of soil thaw was dependent on air temperature and took 5 weeks in 2005, but 10 weeks in 2004. There were two substantial frost events during the recovery period in 2004 that may explain the extended recovery period. In 2005, recovery was complete after the accumulation of 200 growing degree days above 0 degrees C after the start of soil thaw.     

Impact of bark wounds on sapwood in Norway spruce and silver fir

Bark wounds by damage during harvesting are a serious problem in forestry due to fungi infection and wood deterioration. This paper presents results of an investigation about the influence of the wounds on the internal structure of such injured stems. In an experiment, bark wounds were artificially created at the stem base of Norway spruce [Picea abies (L.) Karst.] and silver fir (Abies alba Mill.). 2 years later, the injured stems along with undamaged controls were cut and the trunk portion below breast height subjected to computer tomographic (CT) analysis. Analysis of the CT-images revealed a substantial impact of wounding on sapwood properties in spruce: directly adjacent to the wound surface in all examined trees, a large disturbance zone was detected affecting on average 17% of the potential sapwood area. With increasing distance from the wound, the size of this disturbance zone diminished, but was still detectable in all trees at breast height ca. 1 m above the bark wound.     

Changes in the water relations of balsam fir and white birch saplings after thinning

In the autumn of 1987, young balsam fir (Abies balsamea (L.) Mill.) and white birch (Betula papyrifera Marsh.) trees were thinned and their water relations followed during the next two growing seasons. At the beginning of the first summer following treatment, thinned trees of both species had lower osmotic potentials at full saturation (Psi(pi,sat)) and at turgor loss point (Psi(pi,tlp)) compared with controls. At this time, Psi(pi,sat) was linearly related to the percentage of full sunlight reaching the trees. A higher sugar concentration in leaves was an important component of the lower Psi(pi,sat) of thinned trees. For the other two sampling dates during the first growing season after treatment and all three sampling dates during the second growing season after treatment, little osmotic adjustment of the thinned trees relative to the control tress was observed in either species. The absence of osmotic adjustment during the second growing season following thinning suggests that other mechanisms were responsible for the acclimation of the treated trees to the higher atmospheric evaporative demand. Sapwood permeability (k) of white birch was higher than that of balsam fir, but no differences in k or in sapwood area were found between treated and control trees of either species. Predawn water potentials (Psi(pred)) of treated trees were less negative than those of controls.     

Broadleaf competition interferes with balsam fir regeneration following experimental removal of moose

Changes to ecosystems caused by introduced herbivores can be predictable, stepwise transitions or unpredictable and even irreversible state changes. This study's objectives were to explore effects on forest succession and soil development 5 years after moose (Alces alces L.) were fenced out of areas within and adjacent to a national park in Newfoundland, Canada. Study plots spanned a range of understorey broadleaf plant associations with regenerating balsam fir (Abies balsamea (L.) Mill.), an important winter forage plant for moose and a dominant canopy tree throughout Newfoundland. After 5 years, height–diameter ratios were significantly larger for larger basal diameters of understorey balsam fir in unfenced, but not in fenced subplots, suggesting that growth of the conifer is compromised within the exclosure. In contrast, for most broadleaf trees and shrubs, moose removal by fencing results in greater heights and basal diameters than in control subplots. The competitive advantage of broadleaf trees and shrubs over balsam fir in the short-term may be a result of past sustained heavy moose browsing benefiting plants that are better at investing resources into below-ground growth or benefiting plants that have broader leaf canopies. It is not clear how long the broadleaf transition state we document will continue. Restorative actions intended to mimic usual patterns of forest regeneration in this region of Newfoundland might best consider moose removal with site preparation and/or planting to historic densities.     

Climate influences the leaf area/sapwood area ratio in Scots pine

We tested the hypothesis that the leaf area/sapwood area ratio in Scots pine (Pinus sylvestris L.) is influenced by site differences in water vapor pressure deficit of the air (D). Two stands of the same provenance were selected, one in western Scotland and one in eastern England, so that effects resulting from age, genetic variability, density and fertility were minimized. Compared with the Scots pine trees at the cooler and wetter site in Scotland, the trees at the warmer and drier site in England produced less leaf area per unit of conducting sapwood area both at a stem height of 1.3 m and at the base of the live crown, whereas stem permeability was similar at both sites. Also, trees at the drier site had less leaf area per unit branch cross-sectional area at the branch base than trees at the wetter site. For each site, the average values for leaf area, sapwood area and permeability were used, together with values of transpiration rates at different D, to calculate average stem water potential gradients. Changes in the leaf area/sapwood area ratio acted to maintain a similar water potential gradient in the stems of trees at both sites despite climatic differences between the sites.     

Past and present outbreaks of the balsam fir sawfly in western Newfoundland: An analytical review

Historical data of defoliation and population density were examined to determine whether a sustained outbreak of balsam fir sawfly (Neodiprion abietis Harris) in western Newfoundland, Canada is unprecedented in severity and duration. Results indicate that the current outbreak departs substantially from historical trends, covering a surface area twice the sum of all infestations occurring in the preceding 50 years. The current outbreak is also of longer duration due to a northward expansion of the range usually subjected to severe defoliation by this insect. Time-series analysis indicates that balsam fir sawfly dynamics have a strong second-order component, providing testable hypotheses for future studies investigating the factors responsible for population fluctuations.     

Relationships of climate and cell features in stems and roots of black spruce and balsam fir

? The anatomical differences of mature black spruces and balsam firs were examined at stem and root level in order to characterize their wood properties at cellular level and link these differences to climate.

? Anatomical variability of these species was evaluated in relation to climate data gathered from 2001 to 2004 during the cell enlargement (CE) and wall thickening and lignification (WTL) phases. Lumen area, single cell wall thickness and total tracheid radial diameter were analyzed and regrouped into earlywood and latewood.

? Results from a principal component analysis (PCA) indicated that both first eigenvectors account for 82% and 90% of total variance for CE and WTL respectively. These component factors revealed that precipitation, humidity and number of days with precipitation significantly influence the lumen area (p = 0.0168) and radial cell diameter (p = 0.0222) in earlywood. Significant differences were registered between species and tree parts (stem and root) for the lumen area, radial cell diameter and cell wall thickness in both earlywood and latewood.

? In our study, black spruce exhibited smaller tracheid size in both stem and roots compared to balsam fir. Furthermore, the lower amount of tracheids produced during the growing season and higher proportion of latewood ensure a higher wood density of black spruce. The influence of temperature on earlywood formation is significant, whereas no influence was observed on latewood.

     

A non-invasive method for reconstructing the relative mortality rates of trees in mixed-age,mixed-species forests

The death of overstory trees drives changes throughout forest ecosystems. Knowledge of mortality rates for these larger trees provides a long-term perspective on forest development and forest health. Tree mortality rates are typically determined by repeated censusing of trees over extended time intervals. We describe a method of reconstructing relative mortality rates that does not require injuring live trees on the study plots. To determine when trees died, we used cross-dating (matching tree-ring patterns of dead and live trees), identification of sapling releases, and assessment of tree decomposition. Dead trees were identified to genus or sub-genus by microscopic examination of wood. We reconstructed live tree community structure so that mortality could be relativized on a taxonomic-group- and tree-size-specific basis by the tree densities existing when mortality occurred. We modified a forest development model to operate backwards in time and validated this model by comparing past tree stem diameters predicted by the model with past stem diameters known from tree-ring measurements. All tree-ring measurements from live trees were obtained off the study plots. We report data for all trees ≥20 cm diameter at breast height (DBH) from seven oak-hickory forest sites in Arkansas, Illinois, Indiana, and Ohio, USA. Total plot area was 6.7 ha. Mortality rates were determined for the past 20 years by genus or sub-genus, DBH, and decade. The average mortality rate was 0.60% per year. The accuracy of the reconstructed mortality rates approaches that of mortality rates obtained by repeated censusing. While this mortality reconstruction method does not yield data for individual species, and may involve some compromise in accuracy, the method offers substantial benefits: long-term mortality data may be obtained from a short-term study, mortality rates may be obtained for the past, and because this reconstruction method is non-invasive, future mortality rates can be measured on the same plots.     

Effects of a spruce budworm outbreak on element export below the rooting zone: a case study for a balsam fir forest

? Spruce budworm outbreaks are among the major natural disturbances affecting the dynamics and functioning of Canadian boreal forests. However, the element losses potentially associated with spruce budworm outbreaks have not been quantified.

? We evaluated the influence of spruce budworm outbreaks on nutrient export from boreal forest soils by comparing nutrient leaching losses during a spruce budworm outbreaks episode (1981–1984) to an unperturbed period (1999–2003) in a calibrated catchment located in a balsam fir forest.

? Nutrient soil leaching losses were significantly higher during the spruce budworm outbreaks (1981–1984) for N-NO₃ (30.1 fold), K (8.3 fold), N-NH₄ (6.2 fold), Mg (2.7 fold) and SO₄ (2.2 fold), as compared to an unperturbed period (1999–2003). When the recurrence of spruce budworm outbreaks (33 years) and a plausible average length of such events (5 years) are taken into consideration, it is estimated that in the long term, 5.6 more NO₃, 1.5 more K and 1.2 more NH₄ are leached from the soil profile during outbreaks.

? The important leaching losses during spruce budworm outbreaks, when added to the losses due to tree harvesting and fire (and acid deposition for K), may have considerable effects on soil fertility and ecosystem sustainability.

     

Neutralization and buffering capacity of leaves of sugar maple, largetooth aspen, paper birch and balsam fir

We compared the acidity, the external acid neutralizing capacity and the buffering capacity of leaves of four commercially important tree species, largetooth aspen (Populus grandidentata Michx.), sugar maple (Acer saccharum Marsh.), paper birch (Betula papyrifera Marsh.) and balsam fir (Abies balsamea (L.) Mill), at two sites of contrasting soil fertility in southern Quebec. External acid neutralizing capacity (ENC) of leaves was determined by measuring the change in pH induced by soaking fresh leaves in an acidic solution (pH 4.0) for two hours. The ENC was highest for largetooth aspen (14.3 micro equiv H(+) g(-1)), and lowest for sugar maple and balsam fir (< 5 micro equiv H(+) g(-1)). The buffering capacity index (BCI) was determined by measuring the amount of acid necessary to produce a change of 5 micro equiv H(+) in the leaf homogenate. The BCI ranged from 883 micro equiv H(+) g(-1) for largetooth aspen to less than 105 micro equiv H(+) g(-1) for sugar maple and balsam fir. Leaves of sugar maple and balsam fir had a lower internal pH and a higher percentage of ENC over BCI than paper birch and largetooth aspen. Overall, ENC was correlated with the concentration of all leaf nutrients except Ca, and BCI was correlated with Mg, N and Ca. The site effect was relatively unimportant for all variables.