Contrasting distribution and seasonal dynamics of carbohydrate reserves in stem wood of adult ring-porous sessile oak and diffuse-porous beech trees

We tested the hypothesis that broad-leaved forest species with contrasting wood anatomy and hydraulic system (ring-porous versus diffuse-porous) also differ in distribution and seasonal dynamics of carbohydrate reserves in stem wood. Total nonstructural carbohydrate (TNC) reserves (starch and sugars) were measured enzymatically in the 10 youngest stem xylem rings of adult oak (Quercus petraea (Matt.) Liebl.) and beech (Fagus sylvatica L.) trees during an annual cycle. Radial distribution of carbohydrates was investigated according to ring age. On all dates, oak trees had twofold higher TNC concentration than beech trees (41 versus 23 mg g(DM)(-1)), with starch accounting for the high TNC concentration in oak. Seasonal dynamics of TNC concentration were significantly (P < 0.05) more pronounced in oak (20-64 mg TNC g(DM)(-1)) than in beech (17-34 mg TNC g(DM)(-1)). A marked decrease in TNC concentration was observed in oak trees during bud burst and early wood growth, whereas seasonal fluctuations in TNC concentrations in beech trees were small. The radial distribution of TNC based on ring age differed between species: TNC was restricted to the sapwood rings in oak, whereas in beech, it was distributed throughout the wood from the outermost sapwood ring to the pith. Although the high TNC concentrations in the outermost rings accounted for most of the observed seasonal pattern, all of the 10 youngest xylem rings analyzed participated in the seasonal dynamics of TNC in beech trees. The innermost sapwood rings of oak trees had low TNC concentrations. Stem growth and accumulation of carbon reserves occurred concomitantly during the first part of the season, when there was no soil water deficit. When soil water content was depleted, stem growth ceased in both species, whereas TNC accumulation was negligibly affected and continued until leaf fall. The contrasting dynamics and distribution of carbohydrate reserves in oak and beech are discussed with reference to differences in phenology, early spring growth and hydraulic properties between ring-porous trees and diffuse-porous trees.     

Calibration of thermal dissipation sap flow probes for ring- and diffuse-porous trees

Thermal dissipation probes (the Granier method) are routinely used in forest ecology and water balance studies to estimate whole-tree transpiration. This method utilizes an empirically derived equation to measure sap flux density, which has been reported as independent of wood characteristics. However, errors in calculated sap flux density may occur when large gradients in sap velocity occur along the sensor length or when sensors are inserted into non-conducting wood. These may be conditions routinely associated with ring-porous species, yet there are few cases in which the original calibration has been validated for ring-porous species. We report results from laboratory calibration measurements conducted on excised stems of four ring-porous species and two diffuse-porous species. Our calibration results for ring-porous species were considerably different compared with the original calibration equation. Calibration equation coefficients obtained in this study differed by as much as two to almost three orders of magnitude when compared with the original equation of Granier. Coefficients also differed between ring-porous species across all pressure gradient conditions considered; however, no differences between calibration slopes were observed for data collected within the range of expected in situ pressure gradients. In addition, dye perfusions showed that in three of the four ring-porous species considered, active sapwood was limited to the outermost growth ring. In contrast, our calibration results for diffuse-porous species showed generally good agreement with the empirically derived Granier calibration, and dye perfusions showed that active sapwood was associated with many annual growth rings. Our results suggest that the original calibration of Granier is not universally applicable to all species and xylem types and that previous estimates of absolute rates of water use for ring-porous species obtained using the original calibration coefficients may be associated with substantial error.     

Statics and kinetics of water vapor sorption of small loblolly pine samples

A better understanding of the sorption behavior of different wood structures could be useful in protecting wood against wood deterioration and fungal attack. The purpose of this study was to investigate the effect of differences among earlywood, latewood, and tree ring location within the stem cross-section of loblolly pine (Pinus taeda) on the sorption kinetics and statics of water vapor under ambient conditions. The water vapor sorption of earlywood and latewood in different tree rings was recorded using a dynamic contact angle analyzer under relative humidity changes from 11 to 89%, as provided by saturated salt solutions. Earlywood had higher sorption rates and diffusion coefficients than latewood, while outer tree rings had higher sorption rates and diffusion coefficients than inner tree rings. The sorption isotherms of earlywood, latewood, and different tree ring locations within the stem cross-section were fitted very well by a Hailwood–Horrobin model.     

Refined pipe theory for mechanistic modeling of wood development

We present a mechanistic model of wood tissue development in response to changes in competition, management and climate. The model is based on a refinement of the pipe theory, where the constant ratio between sapwood and leaf area (pipe theory) is replaced by a ratio between pipe conductivity and leaf area. Simulated pipe conductivity changes with age, stand density and climate in response to changes in allocation or pipe radius, or both. The central equation of the model, which calculates the ratio of carbon (C) allocated to leaves and pipes, can be parameterized to describe the contrasting stem conductivity behavior of different tree species: from constant stem conductivity (functional homeostasis hypothesis) to height-related reduction in stem conductivity with age (hydraulic limitation hypothesis). The model simulates the daily growth of pipes (vessels or tracheids), fibers and parenchyma as well as vessel size and simulates the wood density profile and the earlywood to latewood ratio from these data. Initial runs indicate the model yields realistic seasonal changes in pipe radius (decreasing pipe radius from spring to autumn) and wood density, as well as realistic differences associated with the competitive status of trees (denser wood in suppressed trees).     

植物激素与木材形成

综述了植物激素在树木维管组织分化中的作用．生长素是调节木质部细胞分化的主要因子；细胞分裂素在诱导分生组织发端和分生细胞分化上起重要作用；赤霉素促进纤维的形成，赤霉素和生长素结合使用，可有效地促进次生木质部中纤维分化；乙烯能促进树木韧皮部和木质部分化。还讨沦了木材形成过程中的三个主要问题：生长素含量的多少直接控制了沿树轴木质部导管、管胞的大小和密度；外界环境因子影响木质部细胞分化；温带落叶阔叶树环孔材和散孔材形成的机理，生长素控制环孔材树种早材宽导管的形成以及晚材窄导管和纤维形成的假说。对今后改善树木木材的质和量上有一定参考价值。     

A Discussion on Drought Tolerance Through Hydraulic Architecture of Woody Plants

In this paper, ten woody plants grew on the campus of the Beijing Forestry University were selected for measuring hydraulic architecture parameters and water potential of one-year-old twigs. The results show that day and night change of water potential and hydraulic architecture parameters appear to be obviously convex trend. The relationship models between water potential and hydraulic conductivity, special conductivity and leaf special conductivity were respectively established, which were simulated by equations Y=axb and Y=ax2 bx c, through which the parameters physiological meanings were discussed. The results show also show that embolism vulnerability of ten tree species was Robinia pseudoacacia> Gleditsia sinensis> Acer truncatum> Prunus davidiana> Cornus alba> Platycladus orientalis> Cedrus deodara> Pinus bungeana> Pinus tabulaeformis> Jasminum nudiflorum, and that the water transport efficiency in xylem and embolism vulnerability of the species are ring-porous species> diffuse-porous species>non     

Wood specific gravity-mechanical property relationship at species level

Summary Based on the data set of specimen tests on 16 timber species belonging to four distinct wood categories, the specific gravity-mechanical property relationship at species level was examined, and differences in the relationship between species from distinct wood categories were discussed. The linear equation (S =a +bG) was compared with the curvilinear one (S = G) in terms of the goodness at predicting mechanical properties through specific gravity at species level. The specific gravity-mechanical property relationship, to a differing extent, varies with mechanical properties and wood categories. Among three mechanical properties studied, MOR is most closely and almost linearly related to specific gravity, followed by Cmax, whereas MOE is poorly and least linearly related to specific gravity. In general, the relationship between MOE and specific gravity in a species from the ring-porous category is stronger than in a species from the diffuse-porous category. It appears that Cmax in a species from the second softwood category and the ring-porous category is more closely related to specific gravity than in a species from the first softwood category and the diffuse-porous category, respectively. In addition, MOE in a softwood species is generally less related to specific gravity as compared to a hardwood species. Yet, Cmax in a softwood species appears more closely related to specific gravity. Overall, the curvilinear equation is better than the linear one at predicting mechanical properties (especially MOE) in a species.     

A case-study of water transport in co-occurring ring- versus diffuse-porous trees: contrasts in water-status, conducting capacity, cavitation and vessel refilling

Recent work has suggested that the large earlywood vessels of ring-porous trees can be extraordinarily vulnerable to cavitation making it necessary that these trees maintain a consistent and favorable water status. We compared cavitation resistance, vessel refilling, transport capacity and water status in a study of ring-porous Quercus gambelii Nutt. (oak) and diffuse-porous Acer grandidentatum Nutt. (maple). These species co-dominate summer-dry foothills in the western Rocky Mountains of the USA. Native embolism measurements, dye perfusions and balance pressure exudation patterns indicated that the large earlywood vessels of 2-3-year-old oak stems cavitated extensively on a daily basis as predicted from laboratory vulnerability curves, resulting in a more than 80% reduction in hydraulic conductivity. Maple branches showed virtually no cavitation. Oak vessels refilled on a daily basis, despite negative xylem pressure in the transpiration stream, indicating active pressurization of embo-lized vessels. Conductivity and whole-tree water use in oak were between about one-half and two-thirds that in maple on a stem-area basis; but were similar or greater on a leaf-area basis. Oak maintained steady and modest negative xylem pressure potentials during the growing season despite little rainfall, indicating isohydric water status and reliance on deep soil water. Maple was markedly anisohydric and developed more negative pressure potentials during drought, suggesting use of shallower soil water. Although ring porosity may have evolved as a mechanism for coping with winter freezing, this study suggests that it also has major consequences for xylem function during the growing season.     

Genetic control of intra-ring wood density variation in hybrid larch (Larix gmelinii var. japonica × L. kaempferi) F1

Genetic parameters for various wood density traits were estimated in 29-year-old trees of 18 full-sib families of hybrid larch (Larix gmelinii var. japonica × Larix kaempferi) F₁. Intra-ring density variation (IDV) was also evaluated using a model that expresses the pattern curve from earlywood to latewood as a power function. A high IDV indicates an abrupt change in wood density from earlywood to latewood. The ring width and wood density traits of individual rings were determined by X-ray densitometry. Overall wood density (RD) was shown to increase with increasing ring number, ranging from 0.42–0.59 g/cm³, whereas IDV of individual rings decreased gradually from pith outwards. Estimates of individual tree narrow-sense heritability of RD and IDV were 0.66 and 0.67, respectively. IDV showed negative genetic and phenotypic correlations with RD (r _g = −0.99, r _p = −0.72). The predicted genetic gains in latewood proportion and IDV were higher than that of RD. These results suggest that the intra-ring density variation is under moderate genetic control equivalent to wood density. The trend of increasing wood density from earlywood to latewood was associated with changes in both tracheid diameter and cell wall thickness.     

Scaling of angiosperm xylem structure with safety and efficiency

We tested the hypothesis that greater cavitation resistance correlates with less total inter-vessel pit area per vessel (the pit area hypothesis) and evaluated a trade-off between cavitation safety and transport efficiency. Fourteen species of diverse growth form (vine, ring- and diffuse-porous tree, shrub) and family affinity were added to published data predominately from the Rosaceae (29 species total). Two types of vulnerability-to-cavitation curves were found. Ring-porous trees and vines showed an abrupt drop in hydraulic conductivity with increasing negative pressure, whereas hydraulic conductivity in diffuse-porous species generally decreased gradually. The ring-porous type curve was not an artifact of the centrifuge method because it was obtained also with the air-injection technique. A safety versus efficiency trade-off was evident when curves were compared across species: for a given pressure, there was a limited range of optimal vulnerability curves. The pit area hypothesis was supported by a strong relationship (r2 = 0.77) between increasing cavitation resistance and diminishing pit membrane area per vessel (A(P)). Small A(P) was associated with small vessel surface area and hence narrow vessel diameter (D) and short vessel length (L)--consistent with an increase in vessel flow resistance with cavitation resistance. This trade-off was amplified at the tissue level by an increase in xylem/vessel area ratio with cavitation resistance. Ring-porous species were more efficient than diffuse-porous species on a vessel basis but not on a xylem basis owing to higher xylem/vessel area ratios in ring-porous anatomy. Across four orders of magnitude, lumen and end-wall resistivities maintained a relatively tight proportionality with a near-optimal mean of 56% of the total vessel resistivity residing in the end-wall. This was consistent with an underlying scaling of L to D(3/2) across species. Pit flow resistance did not increase with cavitation safety, suggesting that cavitation pressure was not related to mean pit membrane porosity.     

Influence of growth rate on specific gravity and other selected properties of loblolly pine

Summary Forty lobiolly pine trees were studied for the purpose of finding a possible reason for the growth rate-specific gravity controversy. In addition, properties such as specific gravity of earlywood and latewood, latewood percentage, and their relations with growth rate and specific gravity of wood were investigated.The results show that, in general, significant relationship do exist between specific gravity and number of rings per inch for growth rates ranging from approximately 3...8 rings per inch. Specific gravity was generally unrelated with slower growth rates, i.e., more than 8 rings per inch.Specific gravity of specimens having the same number of rings per inch varied according to height-decreasing with increasing height. Latewood percentage was related to number of rings per inch in a similar manner as specific gravity. Within a tree, latewood percentage of wood having the same number of rings per inch decreased with increasing height.Negative relationships between earlywood specific gravity and number of rings per inch were found to be significant for the 30 percent height and for all heights when combined. However, there was no significant relationship between latewood specific gravity and number of rings per inch. Average laterwood specific gravity decreased with height. Earlywood specific gravity only decreased from 5.3...30 percent height levels and then showed an increase. In general, wood properties such as specific gravity and latewood percentage of wood, specific gravity of earlywood and latewood, varied more from breast height to approximately the 30 percent height, than between equal distances above the 30 percent height level.     

The anatomical traits of trunk wood and their relevance to oak (Quercus robur L.) vitality

Oaks’ decline in vitality is attributed to a complex process that involves interactions of several factors leading to increased trees’ mortality. This study investigates the structure of trunk wood of oaks with reference to its physiological role in hydraulic conductivity. On the basis of the crown condition, the oaks were classified into three health groups: healthy trees, declining trees and dead trees. Anatomical traits of wood, such as annual ring width, vessel density, vessel diameter of earlywood and theoretical hydraulic conductivity, were measured and calculated. The narrowest annual rings formed by the cambium were observed in dead oaks. These trees were also characterized by the smallest diameter of earlywood vessels, not only in the period of occurrence of dieback symptoms, but also during their whole life. It is suggested that the formation of narrow annual rings and earlywood vessels of small diameter increases susceptibility of a tree to decay. A reduced vessel diameter implies changes in hydraulic conductivity of oak trunks and thus impairs the water transport, which affects the health of trees. The process of oak decline is considered to have characteristics of natural selection and leads to the elimination of the weakest trees.     

Variability of selected wood characteristics in 40 half-sib families of black spruce (Picea mariana)

Summary Based on 15-year-old black spruce (Picea mariana) trees from 40 half-sib families sampled from 9 blocks of a family test in New Brunswick, this study examined intertree and intratree variation in various wood density and ring width characteristics. Of various variance components of the intertree variation, a remarkable variance component due to family was found in wood density characteristics (viz. average wood density, average earlywood density and latewood density of the tree), and these characteristics are thus under strong genetic control (h _i ² ranging from 0.60 to 0.86, and h _f ² from 0.56 to 0.68). It, to a lesser extent, applies to ring width characteristics at the tree level (viz. average ring width, and average earlywood width, latewood width and latewood percent of the tree) that show a lower heritability (h? from 0.18 to 0.28, and h _f ² from 0.22 to 0.36). Both block and block × family interaction contribute little to the total intertree variation encountered in 40 families from 9 blocks, while tree-to-tree variation within the family accounts for most (over 3/4) of the total intertree variation.Compared to the intertree variation (tree-to-tree variation within the family), the intratree variation in various wood characteristics studied is considerably larger in this species. It appears that most intraring wood density characteristics show a relatively smaller intertree variation but a relatively larger intratree variation as compared to ring width characteristics (except latewood width and latewood percent). Latewood width and latewood percent show the smallest intertree variation and the largest intratree variation. Between the two sources of the radial intratree variation, cambial age explains much more variation in most intraring wood density characteristics, while ring width accounts for more variation in earlywood width, latewood width and intraring density variation. This indicates that wood density of growth rings in this species is dependent more on cambial age than ring width (growth rate). Among various wood density and ring width characteristics studies, maximum (latewood) density shows the strongest response to calendar year. This characteristics is thus a useful dendroclimatic parameter in this species.I would like to thank Dr. E.K. Morgenstern and Mr. D. Simpson for their involvement in the planning of this study. Thanks are also due to G. Chauret, T. Keenam, R. Ploure, V. Steel and C. Reitlingshoefer for their technical assistance     

Effects of ring-porous and diffuse-porous stem wood anatomy on the hydraulic parameters used in a water flow and storage model

Calibration of a recently developed water flow and storage model based on experimental data for a young diffuse-porous beech tree (Fagus sylvatica L.) and a young ring-porous oak tree (Quercus robur L.) revealed that differences in stem wood anatomy between species strongly affect the calibrated values of the hydraulic model parameters. The hydraulic capacitance (C) of the stem storage tissue was higher in oak than in beech (939.8 versus 212.3 mg MPa(-1)). Model simulation of the elastic modulus (epsilon) revealed that this difference was linked to the higher elasticity of the stem storage tissue of oak compared with beech. Furthermore, the hydraulic resistance (R (x)) of beech was about twice that of oak (0.1829 versus 0.1072 MPa s mg(-1)). To determine the physiological meaning of the R (x) parameter identified by model calibration, we analyzed the stem wood anatomy of the beech and oak trees. Calculation of stem specific hydraulic conductivity (k (s)) of beech and oak with the Hagen-Poiseuille equation confirmed the differences in R (x) predicted by the model. The contributions of different vessel diameter classes to the total hydraulic conductivity of the xylem were calculated. As expected, the few big vessels contributed much more to total conductivity than the many small vessels. Compared with beech, the larger vessels of oak resulted in a higher k (s) (10.66 versus 4.90 kg m(-1) s(-1) MPa(-1)). The calculated ratio of k (s) of oak to beech was 2, confirming the R (x) ratio obtained by model calibration. Thus, validation of the R (x) parameter of the model led to identification of its physiological meaning.     

Climatic signal of earlywood vessels of oak on a maritime site

Earlywood vessel lumen areas were measured in 72 consecutive tree rings in wood cores from oak (Quercus robur L.) trees in a maritime woodland. This anatomical time series was statistically correlated with climate data for the same time span. There was a strong dependence of earlywood vessel lumen area on rainfall between February and April, which reflects the role of water availability in vessel ontogeny. By inversion, earlywood vessel lumen areas can be used as a proxy to reconstruct spring precipitation beyond the archived weather records. Such information may be of value in the context of climate change.     

木腐菌及其腐朽机理研究进展

木腐菌可以有效地改善针叶材心材的渗透性,可用于辅助阔叶材散孔材年轮的鉴定,木腐菌及其所分泌的酶还可作为生物技术应用于木材工业,如生物制浆预处理、生物漂白以及木材酶解为糖的预处理等领域。本文从褐腐、白腐、软腐、竹材及人造板等几个方面出发,对国内外腐朽机理的研究现状进行了较为系统、全面的介绍,旨在使学者了解真菌选择性地降解木竹材的化学成分和细胞各部分的知识,为木竹材的保护和合理利用提供参考。     

Morphological changes of Japanese beech treated with the ionic liquid, 1-ethyl-3-methylimidazolium chloride

The morphological changes in wood tissues of Japanese beech (Fagus crenata) upon treatment with the ionic liquid, 1-ethyl-3-methylimidazolium chloride ([C2mim][Cl]), which can dissolve cellulose, were investigated. Treatment with [C2mim][Cl] induced significant swelling of all wood tissues. However, the swelling behavior of wood fibers was different from that of vessels. Intervascular pits were occluded, and pit membranes in ray-vessel pits were broken after treatment with [C2mim][Cl]. No significant differences in swelling behavior were found between latewood and earlywood, although different morphological changes for latewood and earlywood during [C2mim][Cl] treatment were seen in our previous studies on sugi (Cryptomeria japonica). We have found that the effects of [C2mim][Cl] on Japanese beech tissues are inhomogeneous and different from those found for other wood species.     

Pulse-labelling trees to study carbon allocation dynamics: a review of methods, current knowledge and future prospects

Pulse-labelling of trees with stable or radioactive carbon (C) isotopes offers the unique opportunity to trace the fate of labelled CO(2) into the tree and its release to the soil and the atmosphere. Thus, pulse-labelling enables the quantification of C partitioning in forests and the assessment of the role of partitioning in tree growth, resource acquisition and C sequestration. However, this is associated with challenges as regards the choice of a tracer, the methods of tracing labelled C in tree and soil compartments and the quantitative analysis of C dynamics. Based on data from 47 studies, the rate of transfer differs between broadleaved and coniferous species and decreases as temperature and soil water content decrease. Labelled C is rapidly transferred belowground-within a few days or less-and this transfer is slowed down by drought. Half-lives of labelled C in phloem sap (transfer pool) and in mature leaves (source organs) are short, while those of sink organs (growing tissues, seasonal storage) are longer. (13)C measurements in respiratory efflux at high temporal resolution provide the best estimate of the mean residence times of C in respiratory substrate pools, and the best basis for compartmental modelling. Seasonal C dynamics and allocation patterns indicate that sink strength variations are important drivers for C fluxes. We propose a conceptual model for temperate and boreal trees, which considers the use of recently assimilated C versus stored C. We recommend best practices for designing and analysing pulse-labelling experiments, and identify several topics which we consider of prime importance for future research on C allocation in trees: (i) whole-tree C source-sink relations, (ii) C allocation to secondary metabolism, (iii) responses to environmental change, (iv) effects of seasonality versus phenology in and across biomes, and (v) carbon-nitrogen interactions. Substantial progress is expected from emerging technologies, but the largest challenge remains to carry out in situ whole-tree labelling experiments on mature trees to improve our understanding of the environmental and physiological controls on C allocation.     

Comparing the intra-annual wood formation of three European species (Fagus sylvatica, Quercus petraea and Pinus sylvestris) as related to leaf phenology and non-structural carbohydrate dynamics

Monitoring cambial phenology and intra-annual growth dynamics is a useful approach for characterizing the tree growth response to climate change. However, there have been few reports concerning intra-annual wood formation in lowland temperate forests with high time resolution, especially for the comparison between deciduous and coniferous species. The main objective of this study was to determine how the timing, duration and rate of radial growth change between species as related to leaf phenology and the dynamics of non-structural carbohydrates (NSC) under the same climatic conditions. We studied two deciduous species, Fagus sylvatica L. and Quercus petraea (Matt.) Liebl., and an evergreen conifer, Pinus sylvestris L. During the 2009 growing season, we weekly monitored (i) the stem radial increment using dendrometers, (ii) the xylem growth using microcoring and (iii) the leaf phenology from direct observations of the tree crowns. The NSC content was also measured in the eight last rings of the stem cores in April, June and August 2009. The leaf phenology, NSC storage and intra-annual growth were clearly different between species, highlighting their contrasting carbon allocation. Beech growth began just after budburst, with a maximal growth rate when the leaves were mature and variations in the NSC content were low. Thus, beech radial growth seemed highly dependent on leaf photosynthesis. For oak, earlywood quickly developed before budburst, which probably led to the starch decrease quantified in the stem from April to June. For pine, growth began before the needles unfolding and the lack of NSC decrease during the growing season suggested that the substrates for radial growth were new assimilates of the needles from the previous year. Only for oak, the pattern determined from the intra-annual growth measured using microcoring differed from the pattern determined from dendrometer data. For all species, the ring width was significantly influenced by growth duration and not by growth rate, which differs from previous studies. The observed between-species difference at the intra-annual scale is key information for anticipating suitability of future species in temperate forests.     

Rapid carbon accumulation within an unmanaged,mixed, temperate woodland

Forest carbon stocks have increased in both Europe and North America in recent decades. National forest inventories are often used to indicate recent carbon dynamics, but the data from unmanaged forests are often incomplete. Here we calculate changing biomass carbon stocks for a mixed, unmanaged British woodland with two different management histories: (1) older growth stands untouched since 1902 and (2) younger growth stands clear felled in 1943 but have developed naturally since. Transects in the older growth have been monitored since 1945 and the younger growth since 1977. Separate estimates of tree carbon (C), soil C and dead wood C were obtained to verify how C is apportioned in these stands. Tree biomass C stocks had approximately doubled in the older growth stands since 1945 and 60% of C was stored in tree biomass, 38% was stored in soil and 2% stored in coarse woody debris. This study suggests that natural older growth stands are storing more C than typical managed forests, with tree biomass the most important compartment for C stores. If management is to be shifted from biomass production to increased C stores, due consideration should be given to the role of unmanaged, older growth forests.