Bole Surface Area Research Articles

Whether respiration in trees can scale isometrically with bole surface area: A test of hypothesis

Respiration measurements of whole tree have been reported to give evidence that the relative respiration per volume/mass unit decreases with increase in tree size. Based on published datasets, a hypothesis that the relative respiration per area unit in trees can be independent of tree surface area was tested. There is a gap in the published data when the allometric studies of tree size do not intercept with studies on tree respiration. The indirect comparison showed that the scaling exponents, volume vs. surface area and respiration vs. stem volume, are slightly larger than they should be for the hypothesis to hold. The data studied showed that the relative respiration per area unit slightly increases with increase in tree surface area. Possible explanations of the relationship include a different distribution of metabolically active parts of stem and higher nitrogen content in larger trees. Also, the published datasets might include large fast growing trees which had higher growth respiration. To close the data gap, an experiment is required in which the respiration measurements should be performed for the same data as the measurements of scaling between stem volume and surface area.

An application of bole surface growth model: a transitional status of ‘−3/2’ rule

The findings regarding the self-thinning rule in forest stands indicate that the rule may have a sort of ‘asymptotic’ status. It means the admittance of two theses: (1) the slopes of self-thinning trajectories may deviate from −3/2 value; (2) in the space ‘size–density,’ a limiting line exists that serves as a ‘goal’ for population dynamics which this dynamics tends to. An application of the simple geometrical model to the Douglas-fir and Scots pine data suggests that the slope of the self-thinning curve will not remain constant during the course of growth and self-thinning of a single forest stand. Most probably, at the initial stages of stand growth, the slope will be less than −3/2, and at old ages of the stand, the slope will be higher than −3/2. The slope −3/2 is thus an obligatory state in the course of self-thinning of a forest stand. At the very time of −3/2 slope, two particular features fit together. One is that the total bole surface area remains constant. Another feature of the −3/2 slope time is that geometric similarity holds in the growth of the forest stand, which is not in a contradiction with the ‘−3/2’ rule as it had been formulated by its authors. That is, the slope −3/2 (1) is a very specific and obligatory state in the process of forest stand growth and (2) is not an asymptote-like ‘goal’ but rather a transitional point (or may be a span) in the time of growth. These two assertions may be called a transitional status of the ‘−3/2’ rule.

A simple theory to link bole surface area, stem density and average tree dimensions in a forest stand

A geometrical model of a forest stand has been analyzed. A forest stand has been modeled as a population of cones which was described by the change of total bole surface area with density $$\hat{S}(N)$$ , relation between density and a horizontal dimension (radius r) r(N), and the relation between vertical dimension (generatrix l) and radius l(r). It has been shown that there are close relationships between $$\hat{S}(N)$$ , l(r) and r(N). In case of $$\hat{S}(N) = const$$ , power exponent of l(r) can be predicted from the power exponent of r(N) and vice versa. A comparison of the model analysis with the data available on Scots pine (Pinus sylvestris L.) stands has been performed. In spite of the model simplicity, its inferences proved to be workable in many cases where the data can be interpreted as a dynamics of an even-aged forest stand. In particular, if the estimation of total bole surface area is constant, the power exponent in the relation of diameter and stand density DBH(SD) can be calculated on the basis of the power exponent in the relation of height and diameter H(DBH) and vice versa. Possible limitations and the meaning of the analysis are discussed.

Estimates of species- and ecosystem-level respiration of woody stems along an elevational gradient in the Luquillo Mountains, Puerto Rico

We measured CO 2 efflux from stems of seven subtropical tree species situated along an elevational gradient in the Luquillo Mountains, Puerto Rico and scaled these measurements up to the landscape level based on modeled and empirical relations. The most important determinants of ecosystem stem respiration were species composition and stem temperature. At a species scale, measured CO 2 efflux per unit bole surface area at a given temperature was highest in the early successional species Cecropia schreberiana and lowest in species that inhabit high elevations such as Micropholis garciniifolia and Tabebuia rigida. Carbon dioxide efflux rates followed a diel pattern that lagged approximately 6 h behind changes in sapwood temperatures. At an ecosystem scale, our simulation model indicates a decreasing trend of stem respiration rates with increasing elevation due to shifts in species composition, lower temperatures and reductions in branch surface area. The highest estimated stem respiration rates were present in the lowland tabonuco forest type and the lowest rates were present in the elfin forest type (mean 7.4 and 2.1 Mg C ha −1 yr −1, respectively). There was slight temperature-induced seasonal variation in simulated stem respiration rates at low elevations, with a maximum difference of 19% between the months of February and July. Our results coincide well with those of Odum and Jordan [Odum, H.T., Jordan, C.F., 1970. Metabolism and evapotranspiration of the lower forest in a giant plastic cylinder. In: Odum, H.T., Pigeon, R.F. (Eds.), A Tropical Rain Forest: A Study of Irradiation and Ecology at El Verde, Puerto Rico. U.S. Atomic Energy Commission, Oak Ridge, TN, pp. I165–I189] for the tabonuco forest type and extend their work by presenting estimates and spatial patterns of woody tissue respiration for the entire mountain rather than for a single forested plot.

Branch surface area and its vertical distribution in coastal Douglas-fir

Wood area index (WAI; total surface area of branches and bole per unit of land area) is an important yet often neglected forest structural attribute. Branchwood surface area, in particular, has significant implications for many ecophysiological processes including total respiration and interception of radiation and rainfall. Branch surface area was estimated at the branch-, tree-, and stand-level for 33 Douglas-fir (Pseduotsuga menziesii [Mirb.] Franco) plantations in the Oregon Coast Range. Patterns in WAI, leaf area index (LAI; total surface area of needles per unit of land area), tree area index (TAI=WAI+LAI) and various ratios of these dimensions were then investigated. The main axes of primary branches (those attached to the main stem) comprised 82±13% of total branchwood surface area. Tree surface area (needles + woody tissue) increased with increasing tree size and crown length, and decreased with greater intensity of Swiss needle cast (SNC). At the stand-level, woody surface area increased with greater stand density and decreased with more severe SNC, but on average it constituted 29±12% of total tree surface area. Branchwood surface area and bole surface area contributed equally to WAI. The variation in WAI for a given LAI has important implications for radiation and rainfall attenuation in these stands and for accurate partitioning of intercepted radiation between photosynthetic and non-photosynthetic tissues.

Leaf Development and Stem Growth of Ash (Fraxinus excelsior) as Affected by Tree Competitive Status

1. Leaf development and stem growth of ash were studied during year 1993, in an experimental area installed in a mixed natural broadleaved stand, where ash trees, aged 20-24 years, formed part of the dominant layer of the stand. Seventeen trees were selected to represent the range of diameter classes and competitive conditions. 2. For each sample tree, stem girth was measured at several heights above the ground, at the beginning and at the end of the growing season, to evaluate stem volume growth. Crown projection surface area before the growing season and tree height growth were used to estimate a foliar volume. Total leaf dry weight of each tree was obtained by collecting leaves in autumn, and total leaf area was obtained from a sample of leaves. 3. To collect leaves without cutting down the trees, tree crowns were wrapped with nets before leaf fall, from a truck equipped with a nacelle mounted on an articulated arm. Leaves were collected as they fell and accumulated in the bottom of the net. 4. Size relationships were established between foliage biomass and stem and crown dimensions (basal area, crown projection area, foliar volume), between leaf area and foliage biomass, and between bole volume increment and basal area. 5. A basic growth equation was established. This equation shows that consideration of bole surface area is essential for understanding the relationship between annual stem volume increment and foliage biomass. 6. Competition between trees was responsible not only for the differences observed in tree dimensions and growth, but also for the differences in foliage efficiency expressed as bole volume increment per unit of foliage biomass.

Estimation of Bole Surface Area and Bark Volume with Monte Carlo Methods

Many attributes of forest trees can be regarded as integrals of some continuous-dimensional characteristic, and thus their estimation is amenable to variance reduction sampling strategies that have been devised and developed in the simulation field. For the estimation of tree characteristics, these methods provide alternatives to the usual estimation by a fitted regression, and in addition, they provide a flexibility to estimate the integral between any two heights on the tree bole. The application of these methods to the estimation of bole surface area and bark volume is explained, and practical aspects of implementation are discussed.

Marking the Southern Pine Beetle (Coleoptera: Scolytidae) with Rubidium within Loblolly Pine for Dispersal Studies

Studies were conducted to improve techniques for the use of elemental marking in dispersal studies of the southern pine beetle, Dendroctonus frontalis Zimmermann. Tissues from host loblolly pines, Pinus taeda L., having diameters at breast height ranging from 15 to 46 cm and the emerging D. frontalis were marked successfully by injection of rubidium (Rb) concentrations based on estimates of bole surface area. A stem-well infusion technique, which is less labor intensive than pressure injections, was found to mark emerging beetles successfully. Use of dimethysulfoxide (DMSO) in RbCI solution did not significantly increase Rb concentration in P. taeda phloem samples or percentage of Rb-marked D. frontalis. Rb was rapidly eliminated by Rb-marked D. frontalis adults when allowed to feed on nonRb-enhanced host tissue, but Rb concentrations remained relatively constant in Rb-marked beetles held without feeding. While this loss of Rb by colonizing D. frontalis would effect use of the elemental labeling technique for subsequent tracking of reemerging adults, it would be relatively unimportant for typical short duration mark-recapture experiments. Rb-marked D. frontalis were successfully recaptured in pheromone-baited traps at distances as far as 1 km from the point of emergence. Recaptures in four cardinal directions indicated a random dispersal pattern. Adirect comparison of fluorescent powder and Rb-marking showed both are equally effective. Selection of method will depend on the objectives and experimental design.

Components of woody-tissue respiration in young Abies amabilis (Dougl.) Forbes trees

Woody-tissue respiration was measured on five different dates at three to five locations on each of 12 30-year-old Abies amabilis trees. On any given date, temperature-corrected respiration per unit surface area varied 10 to 40-fold between sampling locations. In stems, the two major components of respiration were growth respiration and sapwood maintenance respiration, which were of roughly equal importance during the growing season. There was no evidence of significant cambial maintenance respiration, suggesting that a stand with high bole surface area would not automatically have high respiration. Respiration in branches was much greater than in boles of comparable volume and growth rates, and was significantly correlated with branch height. Branch respiration may include an another significant component in addition to the two seen in bole respiration, possibly associated with carbohydrate mobilization and transport or with CO2 efflux from the transpiration stream.

Mass and nutrient content of decaying boles in an Engelmann spruce – subalpine fir forest, Rocky Mountain National Park, Colorado

We classified dead bole wood in an old-growth Engelmann spruce – subalpine fir (Piceaengelmannii Parry – Abieslasiocarpa (Hook.) Nutt.) forest in Rocky Mountain National Park, Colorado, into decay classes and measured dead bole surface area, volume, biomass, and nutrient content. Biomass of dead boles was 70 Mg/ha, about half as large as aboveground live biomass in these forests. Net accumulation of N, P, Ca, and Na occurred with increasing decay. The N:P ratio varied little with decay, approaching a value of 20 in the most decayed wood, typical of that found in other studies of dead boles. Loss of K during bole decay exceeded the rate of weight loss, whereas Mg loss followed weight loss. The total pools of nutrients in dead boles and in parentheses, the amount of nutrients stored in dead boles as a percentage of total above- and below-ground living, forest floor, and dead wood nutrients were 92.2 kg N/ha (7%), 4.89 kg P/ha (5%), 67.9 kg K/ha (16%), 156.6 kg Ca/ha (12%), 28.9 kg Mg/ha (17%), and 0.74 kg Na/ha (9%). Storage of relatively high amounts of Ca in dead wood of most natural forests indicates that management could have a significant effect on its availability in the long term.

Growth and maintenance respiration in stems of Pinuscontorta and Piceaengelmannii

Stem maintenance respiration was linearly related to live-cell volume for lodgepole pine (Pinuscontorta var. latifolia Engelm.) from 4 to 36 cm dbh and for Engelmann spruce (Piceaengelmannii Parry) from 0 to 20 cm dbh. Sapwood contained greater than 80% of the total live-cell volume in stems. Bole surface area, commonly used to estimate tree respiration costs, poorly estimated stem maintenance respiration. At 15 °C, maintenance costs for lodgepole pine were 6.6 × 10−5 kg C•(kg C sapwood)−1•d−1. Stem respiration during the growing season, both corrected and uncorrected for maintenance, correlated well with annual stemwood growth. Annual stem maintenance respiration for trees and stands can be estimated using sapwood volume, sapwood temperature, and knowledge of respiratory behavior. Total respiration (construction plus maintenance) estimated using stem growth and a model of maintenance respiration was compared with actual respiration measurements integrated over a 100-d growing season. Estimated respiration agreed with the integrated measurements for Engelmann spruce, but overestimated the integrated measurements by 73% in lodgepole pine. These results suggest that estimates of stem respiration made during the growing season may be affected by transpiration.

ESTIMATING ATTACK AND BROOD TOTALS AND DENSITIES OF THE MOUNTAIN PINE BEETLE IN INDIVIDUAL LODGEPOLE PINE TREES

AbstractA method was developed for estimating attack and brood totals of the mountain pine beetle in part or all of the infested bole of individual lodgepole pine trees. The method requires measurement of tree height, tree diameter at 1.37 m, and maximum height of attack, and tallying of brood and attacks on paired 20 cm by 30 cm samples centered at 1.22 m on the bole. Attack and brood totals are estimated in two ways: (1) as a product of the variables (a) total bole surface area, (b) proportion of bole surface area infested, and (c) mean attack/brood density; and (2) regression on the combined variables of tree diameter, infested height, and attack/brood density at 1.22 m on the bole. An approach to developing variance estimates for brood and attack totals is discussed.

Calculating Tree Bole Surface Area for Estimating Populations of the Southern Pine Beetle (Coleoptera: Scolytidae)

A conoidal frustum model was determined to be a practical and an accurate method for estimating the bole surface area of southern pines. Sample percent bias and absolute bias criteria indicated clear superiority over a nonlinear regression model that has been widely applied as a tree geometry model in population dynamics studies of the southern pine beetle, Dendroctonus frontalis Zimmermann. The conoidal frustum estimator should be relied upon in all cases where population density data are to be expanded to the total infested area per tree or to groups of trees, with a minimum of percent bias and absolute bias and without systematic bias trends.

Density, Biomass, Productivity, and Nutrient‐Cycling Changes During Stand Development in Wave‐Regenerated Balsam Fir Forests

Regeneration waves in the high—altitude fir forests of the northeastern United States create a gradient of stands of different ages that are ideal for research on forest developmental processes. Changes in density, biomass, productivity, litterfall, and nutrient content of the vegetation over the course of stand development were documented in a series of these stands on Whiteface Mountain, New York. After a regeneration wave passes through an area and the overstory dies, formerly suppressed seedlings are released, and stand density increases to 10—13 trees/m2 at about age 10 (counting only stems over 25 cm tall). Stand density then begins to decrease, at first slowly but then more rapidly, with a maximum mortality rate of 20—25% yr between years 20 and 30. Thereafter mortality continues, but the rate decreases over time to ≈4% yr at age 55. After about age 20 the relationship between stand age and stand density is p = 0.04 · exp(115/AGE). For stands that have begun to self—thin, the relationship between stand age and stand density and mean aboveground tree mass is log10 m = 3.94 — 1.24 log10p, with a slope significantly different from —1.5. Basal area (measured at 25 cm) increases for the first 15—20 yr of stand development, then remains constant at 60—70 m2/ha for the remainder of the life of the stand. Total aboveground biomass of a mature fir stand is ≈11.8 kg/m2 (1.5 kg/m2 foliage, 1.7 kg/m2 branches, 0.9 kg/m2 bole bark, and 7.7 kg/m2 bole wood). The biomass accumulation rate is highest during the first decade of stand development (320 g · m—2 · yr—1), and declines to ≈115 g · m—2 · yr—1 for a 55—yr—old stand. Foliage biomass increases rapidly during the first 10—15 yr of stand development, then remains constant after crown closure. Branch biomass increases until about age 15, declines for about a decade, then increases again for the rest of the life of the stand. Bole bark increases until about age 30, then stabilizes. Bole wood continues to increase throughout the life of the stand. Aboveground net primary productivity for a mature stand is 960 g · m—2 · yr—1 (foliage, 380 g · m—2 · yr—1; twigs, 150 g · m—2 · yr—1; branch wood and bark, 170 g · m—2 · yr—1; bole bark, 25 g · m—2 · yr—1; and bole wood, 235 g · m—2 · yr—1). Productivity remains generally constant as the stand develops, but in the early stages almost 45% of the total aboveground production is devoted to producing bole wood and bark, while in a mature stand this proportion drops to ≈25%, with the remainder going to foliage and branch wood production. There is thus a shift from production of boles to branches as the stand ages. This may reflect a shift in the primary stresses on the trees, from intraspecific competition in the younger stands to physical environmental factors (e.g., wind abrasion and rime formation) in the older stands. After the initial increase, woody tissue respiration remains approximately constant throughout the life of the stand, as an increase in the size and height of boles is balanced by death of small trees and a decrease in total bole surface area. A mature fir stand contains nitrogen, potassium, magnesium, and calcium in the amounts 47.7, 18.1, 4.6, and 30.4 g/m2, respectively. All these nutrients accumulate rapidly in the early stages of stand development as the stand accumulates nutrient—rich foliage. This is particularly true for nitrogen; a 9—yr—old stand contains half as much nitrogen as a 60—yr—old one. However, the difference between nutrient uptake by the trees and nutrient release in decomposition is greatest somewhat later in stand development, when the pulse of litter accompanying stand die—off has decomposed but the young stand is still taking up significantly greater quantities of nutrients than it is returning in litterfall. Thus if ecosystem nutrient losses are controlled by nutrient accumulation in living biomass, the period of maximum nutrient conservation should occur in the years immediately after disturbance, when the stand is accumulating nutrients most rapidly. If the critical factor is the balance between uptake and release, then minimum losses should occur later. Increased meteorologic inputs of nitrogen and acid in the high—altitude fir forests may have altered the natural nutrient cycles of the preindustrial era.