Wind-induced stresses in cherry trees: evidence against the hypothesis of constant stress levels

We calculated the wind-induced bending moments and stresses generated in the stems of five Prunus serotina conspecifics differing in height and canopy shape and size (based on detailed measurements of stem projected area and location with respect to ground level) to test the hypothesis that wind-loads generate uniform and constant stress levels along the lengths of tree twigs, branches, and trunks. These calculations were performed using five different wind speed profiles to evaluate the relative importance of the shape of wind speed profiles versus the ’geometry’ of tree shape on stem stress distributions and magnitudes. Additionally, we evaluated the effect of absolute tree size and stem taper on wind- induced stresses by scaling the size of smaller conspecifics to the absolute height of the largest of the five trees yet retaining the original stem proportions (i.e., diameter relative to stem length) for each plant. Finally, we also determined how the factor of safety for wind-loading (i.e., the quotient of stem yield stress and wind-load stress) changed as a function of tree size (and, presumably, age). Our results indicate that wind-load stress levels (1) vary along stem length even for the same wind speed profile and the same maximum wind speed; (2) would increase to dangerous levels with increasing tree height if it were not for ontogenetic changes in stem taper and canopy shape that reduce stress intensities to manageable levels; (3) tend to be more dependent on stem taper and canopy shape and size than on the shape of the wind speed profile; and (4) the factor of safety against wind-induced mechanical failure decreases as trees get larger, but varies along the length of large trees such that preferential stem failure is likely and functionally adaptive. We thus (1) reject the hypothesis of constant wind-induced stress levels; (2) support the view that size-dependent changes in stem taper are required to maintain wind-load mechanical reliability; and (3) suggest that certain portions of mature trees are ’designed’ to fail under high winds speeds, thereby reducing drag and the bending moments and stresses experienced by trunks. Received: 24 May 1999 / Accepted: 8 October 1999     

Computing factors of safety against wind-induced tree stem damage

The drag forces, bending moments and stresses acting on stems differing in size and location within the mechanical infrastructure of a large wild cherry (Prunus serotina Ehrh.) tree are estimated and used to calculate the factor of safety against wind-induced mechanical failure based on the mean breaking stress of intact stems and samples of wood drawn from this tree. The drag forces acting on stems are calculated based on stem projected areas and field measurements of wind speed taken within the canopy and along the length of the trunk. The bending moments and stresses resulting from these forces are shown to increase basipetally in a nearly log-log linear fashion toward the base of the tree. The factor of safety, however, varies in a sinusoidal manner such that the most distal stems have the highest factors of safety, whereas stems of intermediate location and portions of the trunk near ground level have equivalent and much lower factors of safety. This pattern of variation is interpreted to indicate that, as a course of normal growth and development, trees similar to the one examined in this study maintain a cadre of stems prone to wind-induced mechanical damage that can reduce the probability of catastrophic tree failure by reducing the drag forces acting on older portions of the tree. Comparisons among real and hypothetical stems with different taper experiencing different vertical wind speed profiles show that geometrically self-similar stems have larger factors of safety than stems tapering according to elastic or stress self-similarity, and that safety factors are less significantly influenced by the 'geometry' of the wind-profile.     

Identification of biomechanical factors involved in stem shape variability between apricot tree varieties

BACKGROUND AND AIMS: Stem shape in angiosperms depends on several growth traits such as elongation direction, amount and position of axillary loads, stem dimensions, wood elasticity, radial growth dynamics and active re-orientation due to tension wood. This paper analyses the relationship between these biomechanical factors and stem shape variability. METHODS: Three apricot tree varieties with contrasting stem shape were studied. Growth and bending dynamics, mechanical properties and amount of tension wood were measured on 40 1-year-old stems of each variety during one growth season. Formulae derived from simple biomechanical models are proposed to quantify the relationship between biomechanical factors and re-orientation of the stems. The effect of biomechanical factors is quantified combining their mechanical sensitivity and their actual variability. RESULTS: Re-orientations happened in three main periods, involving distinct biomechanical phenomena: (a) passive bending due to the increase of shoot and fruit load at the start of the season; (b) passive uprighting at the fall of fruits; (c) active uprighting due tension wood production at the end of the season. Differences between varieties mainly happened during periods (a) and (b). CONCLUSIONS: The main factors causing differences between varieties are the length/diameter and the load/cross-sectional area ratios during period (a). Wood elasticity does not play an important role because of its low inter-variety variability. Differences during period (b) are related to the dynamics of radial growth: varieties with early radial growth bend weakly upward because the new wood layers tend to set them in a bent position. The action of tension wood during period (c) is low when compared with passive phenomena involved in periods (a) and (b).     

Reducing stem bending increases the height growth of tall pines

The hypothesis was tested that upper limits to height growth in trees are the result of the increasing bending moment of trees as they grow in height. The increasing bending moment of tall trees demands increased radial growth at the expense of height growth to maintain mechanical stability. In this study, the bending moment of large lodgepole pine (Pinus contorta Dougl. Ex Loud. var. latifolia Engelm.) was reduced by tethering trees at 10 m height to counter the wind load. Average bending moment of tethered trees was reduced to 38% of control trees. Six years of tethering resulted in a 40% increase in height growth relative to the period before tethering. By contrast, control trees showed decreased height growth in the period after tethering treatment. Average radial growth along the bole, relative to height growth, was reduced in tethered trees. This strongly suggests that mechanical constraints play a crucial role in limiting the height growth of tall trees. Analysis of bending moment and basal area increment at both 10 m and 1.3 m showed that the amount of wood added to the stem was closely related to the bending moment produced at these heights, in both control and tethered trees. The tethering treatment also resulted in an increase in the proportion of latewood at the tethering height, relative to 1.3 m height. For untethered control trees, the ratio of bending stresses at 10 m versus 1.3 m height was close to 1 in both 1998 and 2003, suggesting a uniform stress distribution along the outer surface of the bole.     

Biomechanical design and long-term stability of trees: Morphological and wood traits involved in the balance between weight increase and the gravitropic reaction

Studies on tree biomechanical design usually focus on stem stiffness, resistance to breakage or uprooting, and elastic stability. Here we consider another biomechanical constraint related to the interaction between growth and gravity. Because stems are slender structures and are never perfectly symmetric, the increase in tree mass always causes bending movements. Given the current mechanical design of trees, integration of these movements over time would ultimately lead to a weeping habit unless some gravitropic correction occurs. This correction is achieved by asymmetric internal forces induced during the maturation of new wood.The long-term stability of a growing stem therefore depends on how the gravitropic correction that is generated by diameter growth balances the disturbance due to increasing self weight. General mechanical formulations based on beam theory are proposed to model these phenomena. The rates of disturbance and correction associated with a growth increment are deduced and expressed as a function of elementary traits of stem morphology, cross-section anatomy and wood properties. Evaluation of these traits using previously published data shows that the balance between the correction and the disturbance strongly depends on the efficiency of the gravitropic correction, which depends on the asymmetry of wood maturation strain, eccentric growth, and gradients in wood stiffness. By combining disturbance and correction rates, the gravitropic performance indicates the dynamics of stem bending during growth. It depends on stem biomechanical traits and dimensions. By analyzing dimensional effects, we show that the necessity for gravitropic correction might constrain stem allometric growth in the long-term. This constraint is compared to the requirement for elastic stability, showing that gravitropic performance limits the increase in height of tilted stem and branches. The performance of this function may thus limit the slenderness and lean of stems, and therefore the ability of the tree to capture light in a heterogeneous environment.     

An evaluation of the uniform stress hypothesis based on stem geometry in selected North American conifers

The uniform stress hypothesis of stem formation was evaluated by comparing stem taper of Abies balsamea, Abies lasiocarpa, Picea rubens, Pinus contorta, Pinus elliottii, Pinus palustris, Pinus ponderosa, Pinus taeda, and Pseudotsuga menziesii to the taper expected if stems develop to uniformly distribute bending stress. The comparison was conducted by regressing stem diameter at height h (D_h) against bending moment at h (M_h) using the model D_h=J (M_h)^' where J and ' are fitted coefficients, and testing for '=0.333, the hypothesized value. Twelve curves were fitted with the model. Seven of the fitted values of ' were significantly different from 0.333, but eight of the values were within ᆞ% of 0.333 and eleven values were within ᆣ% of 0.333. Where the fitted value of ' was >15% of 0.333, residuals were biased with height. Fit by relative height, values of ' were within ᆞ% of 0.333 for large portions of these stems. While most of the fitted values of ' support the uniform-stress hypothesis, the values of ' for Pseudotsuga menziesii trees clearly did not. Many of the fitted values of J were inversely related to the modulus of elasticity (E) of green wood reported for these species. With the exception of Pseudotsuga menziesii, growing conditions appeared to account for extraordinary values of J. Increases in J with stem height corresponded with reported decreases in E with height. The covariance between J and E suggests some regulation of bending curvature by adjustments in cross-sectional area. These results suggest that stems taper to maintain a uniform bending curvature and that when E is relatively constant within and among stems, diameter along the stem or across stems can be predicted from bending moment using a simple power function.     

Development and Growth Form of the Neotropical Liana <Emphasis Type="Italic">Croton nuntians</Emphasis>: The Effect of Light and Mode of Attachment on the Biomechanics of the Stem

The neotropical liana Croton nuntians (Euphorbiaceae) can occur in a variety of different growth habits. Juvenile freestanding plants are mechanically stable without support and resemble morphologically young trees or shrubs, whereas adult plants are climbers. Ontogenetic variation of bending and torsion properties of different growth phases are analyzed by measurements of flexural stiffness, structural bending modulus, torsional stiffness and structural torsional modulus. Mechanical and anatomical data show two fundamentally different patterns for juvenile freestanding and adult climbing plants. In freestanding plants, mechanical properties and the contribution of cortex, wood, and pith to the stem cross-section vary only little during ontogeny as is typical for semi-self-supporting plants. In contrast, climbing plants become significantly more flexible during ontogeny, as is characteristic for lianas. This is accompanied by a transition to the formation of a less dense wood type with large diameter vessels and an increasing contribution of flexible tissues (less dense wood and cortex) to the cross-sectional area and the axial second moment of area of the stems. Depending on the environmental conditions, freestanding plants can differ considerably in their appearance due to differences in branching system or stem taper. Therefore the influence of light quantity, measured as percentage of canopy opening, on the mechanical properties and the stem anatomy was tested. Freestanding plants grown with strong shade are significantly more stiff in bending compared with plants grown with a moderate light environment.     

Cellulose and lignin biosynthesis is altered by ozone in wood of hybrid poplar (Populus tremula × alba)

Wood formation in trees is a dynamic process that is strongly affected by environmental factors. However, the impact of ozone on wood is poorly documented. The objective of this study was to assess the effects of ozone on wood formation by focusing on the two major wood components, cellulose and lignin, and analysing any anatomical modifications. Young hybrid poplars (Populus tremula × alba) were cultivated under different ozone concentrations (50, 100, 200, and 300 l l(-1)). As upright poplars usually develop tension wood in a non-set pattern, the trees were bent in order to induce tension wood formation on the upper side of the stem and normal or opposite wood on the lower side. Biosynthesis of cellulose and lignin (enzymes and RNA levels), together with cambial growth, decreased in response to ozone exposure. The cellulose to lignin ratio was reduced, suggesting that cellulose biosynthesis was more affected than that of lignin. Tension wood was generally more altered than opposite wood, especially at the anatomical level. Tension wood may be more susceptible to reduced carbon allocation to the stems under ozone exposure. These results suggested a coordinated regulation of cellulose and lignin deposition to sustain mechanical strength under ozone. The modifications of the cellulose to lignin ratio and wood anatomy could allow the tree to maintain radial growth while minimizing carbon cost.     

Variations in maturation strains and root shape in root systems of Maritime pine (Pinus pinaster Ait.)

 In order to determine if different types of wood were being laid down in the root system of Maritime pine (Pinus pinaster Ait), in response to wind loading, longitudinal residual maturation strains (LRMS), indicating the existence of mechanical stress in developing wood cells, were measured in the trunk and lateral roots. Two age groups of trees (5- and 13-year- old) were compared. LRMS were greater in the trunk and roots of 13-year-old trees than in 5-year-old trees. This phenomenon may be due to increased competition between older trees. LRMS in leeward roots of both age-groups were positive i.e. the wood cells had developed under compression, as also occurs in reaction wood of gymnosperms. As leeward roots are placed under compression during tree sway, an abnormal type of wood may form in the roots in order to counteract the increased stress. In other roots, the strains were negative i.e. the cells had developed under tension, as occurs in normal wood. In the roots of younger trees, LRMS were also positive nearer the stem, thus indicating that wood formation may also be influenced by bending stresses experienced in this zone. In addition to LRMS measurements, radial growth in roots was examined in order to determine the influence of mechanical loading on secondary growth. In older trees, there was a significant increase of 34% in woody growth below the biological centre, compared to that above. This eccentricity is unlike that found in most other tree species, where secondary growth is usually greater on the upper side of the root. However, Maritime pine has a tap root, which will alter the pattern of stress within the root system. Under wind loading, a concentration of mechanical stress will develop at the bases of the stem, lateral roots and tap root. Received: 7 July 1997 / Accepted: 11 December 1997     

Relationships between stem structure and bending strength in Pinus radiata seedlings

Summary Seedlings from nine families of Pinus radiata were grown in a glasshouse under conditions of high and low nitrate nitrogen availability to investigate effects on anatomical and strength characteristics of stems. Families were classified into groups dependent upon their previously determined susceptibility to stem deformation prevalent in plantations established on fertile ex-pasture. Nitrogen treatments significantly affected seedling form in terms of both branch production and stem slenderness. The high N treatment resulted in shorter seedlings, a proportion of which were obviously stunted. Stem strength of seedlings, physically supported throughout the experiment, was assessed as stem lean at harvest as well as the bending strength of the fresh stem at 50% stem height. These two variables were found not to be correlated. Stem lean at harvest was greatest in families known to be susceptible to stem deformation. These families produced stems that were also more slender than families of low susceptibility. Increased stem lean was associated mostly with increased stem slenderness while elasticity was more influenced by pith diameter, stem density and wood radius.     

Size-dependent Allometry of Tree Height, Diameter and Trunk-taper

The allometry of tree height with respect to trunk diameterand the allometry of trunk diameter with respect to distancefrom the top of the tree (i.e. trunk taper) were determinedfor 27 Robinia pseudoacacia trees differing in age and sizegrowing in an open field. The allometric (scaling) exponentfor height was > 1 for small and young trees and decreasedto 2/3 and then 1/2 as tree size and age increased. Similarly,the exponent for taper was > 1 near the tips of young andold trunks and converged onto values of 2/3 and 1/2 toward thebase of mature tree trunks. These observations indicate thata single 'optimal mechanical design principle' (i.e. elastic,stress or geometric self-similarity) neither holds true throughoutthe lifetime of R. pseudoacacia trees, nor does a single designprinciple govern the taper of a trunk throughout its entirelength. Rather, over the course of growth and development, theallometry of R. pseudoacacia tree height and trunk taper progressivelychanges, complying with geometric self-similarity for youngplants (and young portions of old plants) and subsequently givingthe appearance of elastic or stress self-similarity as plants(or portions of plants) get older and therefore larger. Analysesof published (and new) data suggest that the conclusions drawnfor R. pseudoacacia trees are likely to hold true for othertree species because stem growth in diameter is 'indeterminate'whereas growth in overall tree height is asymptotic and thereforeessentially 'determinate'.Copyright 1995, 1999 Academic Press Scaling, woody plants, Robinia pseudoacacia     

Effects of experimental stem burial on radial growth and wood anatomy of pedunculate oak

In dendrogeomorphology, abrupt changes in wood anatomy are frequently used to date the exact year of burial and exposure events. However, few studies have addressed the precision and underlying mechanisms of these changes. In a field experiment, performed in a drift-sand area in the Netherlands, we buried the stems of mature pedunculate oak trees (Quercus robur L.) up to a height of 50 cm and analysed the responses in ring width and earlywood-vessel characteristics, while monitoring the course of temperature above and below the soil surface.After 3 years of stem burial, we found no significant differences in ring width and earlywood-vessel characteristics between control and buried trees both above and below the burial level. Burial however strongly reduced temperature amplitude and the occurrence of sub-zero temperatures around the buried stems. All buried trees formed epitropic roots that grew upward into the new sediment layer, but no adventitious roots were formed on the buried stems. Irrespective of the burial treatments, we found that the mean ring width was largest at the original stem base and lowest at breast height. In contrast, vessel sizes were significantly larger at breast height compared with the stem base. Differences in vessel density barely differed between years and heights.In our field experiment on mature pedunculate oak trees, the burial of stems by 50 cm of drift sand did not induce any local growth suppression or detectable changes in wood anatomy. As wood-anatomical changes in response to burial have previously been reported for trees that had formed adventitious roots, we stress the role of adventitious-root formation as a possible trigger behind the local changes in wood anatomy, reflecting a functional change of a buried stem towards a root. Based on our field experiment, it seems unlikely that years of shallow or moderate burial events (≤50 cm) can be reconstructed using the wood structure of buried stems. As epitropic roots develop quickly after burial, dating such roots may potentially yield better estimates of burial events. Further research on the relation between adventitious root and changes in stem anatomy is needed to ascertain the precision of dating sand-burial events using tree rings.     

The Biomechanics of Tree Fork Design

Two biomechanically different types of tree fork are described: the “compression fork” where the two jointed stems are pressed against each other at the contact face by the action of reaction wood, and the “tension fork” where the two connected stems are bent away from each other by gravity or wind action leading to tensile stresses in the connective zone. It is well known that trees permanently try to improve their own designs by adaptive growth in order to maintain a state of constant mechanical stress at the tree surface. In the case of these two different types of tree fork, adaptive growth also takes different ways in order to avoid high localized stress peaks which could lead to failure of the tree under wind loading. In this paper only the tension fork is assessed with respect to its shape optimization by computer simulation of adaptive growth. It is shown that the tensile fork is shape optimized in a very perfect way in order to avoid any dangerous localized stress peaks (notch stresses) which could lead to failure of the tree.     

The influence of stem guying on radial growth,stem form and internal resin features in radiata pine

Key message

Stem guying to prevent wind-induced swaying of radiata pine trees resulted in significant changes in radial growth, but did not affect the frequency of compression wood or resin features.

Abstract

Mechanical stress resulting from wind forces acting on trees can cause a number of direct and indirect effects ranging from microscopic changes in cambial activity through to stem breakage and uprooting. To better understand these effects on radial stem growth and wood properties, an experiment was established in a 13-year-old radiata pine (Pinus radiata D Don) stand in which 20 trees were guyed to prevent them from swaying. Radial growth was monitored in these trees and 20 matched controls at monthly intervals for 5 years. The trees were then felled and radial growth, resin features and compression wood were assessed on cross-sectional discs taken at fixed locations up the stem. There was a significant reduction in radial growth at breast height (1.4 m above the ground) in the guyed trees, but an increase in growth immediately above the guying point. A total of 277 resin features were observed in the growth rings formed following guying. The overall frequency of such features was related to height within the stem and annual ring number. No effect of stem guying was found on the incidence of compression wood. Interestingly, the distribution of resin features also did not differ between guyed and un-guyed trees. There was no evidence of a link between stem restraint as a result of guying and the incidence of resin features, suggesting that other factors, such as soil moisture may be more influential.     

Nitrogen source effects on rhizosphere pH and nutrient accumulation by Pacific Northwest conifers

Growth responses ofCasuarina cunninghamiana to inoculation withFrankia are described in unsterilized field soils at three sites. At Mt Crawford, South Australia, seedlings of three provenances ofC. cunninghamiana were inoculated with a singleFrankia source just prior to planting out. Forty-four months after planting, inoculation had more than doubled wood production by twoC. cunninghamiana provenances, whilst a third provenance grew poorly and did not respond to inoculation. In Zimbabwe, seedlings of one provenance ofC. cunninghamiana were inoculated in the nursery with one of four differentFrankia strains. In an N deficient soil at Kadoma, three of theseFrankia increased tree height 14 months after planting by between 50% and 70% in comparison to the uninoculated seedlings. The fourthFrankia strain resulted in increased tree height to three times that of the uninoculated controls and up to double that of the other threeFrankia strains. At Gympie, Queensland, Australia, seedlings ofC. cunninghamiana raised open-rooted in a nursery bed were inoculated withFrankia seventeen weeks before planting out. During the 22 months following planting in the field, tree growth was limited by soil P status and there was no response in tree height or stem diameter to inoculation withFrankia or to N fertilizer unless P was applied. In the presence of added P there was a significant response both toFrankia inoculation and to N fertilizer. This positive interaction between P application and N treatment was reflected in wood volumes-inoculated trees and those trees supplied N fertilizer produced 34% and 95% more wood volume than did the uninoculated trees. These results demonstrate the potential to increase the productivity of Casuarina plantings by inoculation withFrankia and by alleviation of P deficiency.     

Intra-specific competition and the radial development of wood density, microfibril angle and modulus of elasticity in plantation-grown Eucalyptus nitens

In eucalypt plantations managed for solid-wood products, radial trends in wood density, microfibril angle (MFA) and stiffness (modulus of elasticity, MoE) are properties of potential commercial importance that can be affected by competition from neighbouring trees. In this study, wood properties at breast height (1.3?m) were studied on radial strips prepared from 12-mm pith-to-bark wood cores taken from 20 trees in a 22-year-old Eucalyptus nitens (Deane and Maiden) Maiden thinning trial in north-eastern Tasmania, Australia. Thinning treatments were applied at age 6?years. Trees were sampled from each of the 200, 400?stems ha^?1 and unthinned control treatments. Intra-specific competition for each sampled tree was estimated using the basal area growth of surrounding trees. SilviScan? technology was used to produce radial profiles of wood density, MFA and MoE. Results indicate a reduction in intra-specific competition through thinning of E. nitens plantations at an early age leads to a transient increase in MFA but has no significant effect on wood density or the intra-annual cycle of wood density. The correlation between the level of intra-specific competition and initial change in MFA following thinning, and a significant relationship between tree shape and mean MFA at breast height suggests the change in MFA is a post-thinning response to increased exposure and wind sway.     

Gibberellin-induced formation of tension wood in angiosperm trees

After gibberellin had been applied to the vertical stems of four species of angiosperm trees for approximately 2 months, we observed eccentric radial growth that was due to the enhanced growth rings on the sides of stems to which gibberellin had been applied. Moreover, the application of gibberellin resulted in the formation of wood fibers in which the thickness of inner layers of cell walls was enhanced. These thickened inner layers of cell walls were unlignified or only slightly lignified. In addition, cellulose microfibrils on the innermost surface of these thickened inner layers of cell walls were oriented parallel or nearly parallel to the longitudinal axis of the fibers. Such thickened inner layers of cell walls had features similar to those of gelatinous layers in the wood fibers of tension wood, which are referred to as gelatinous fibers. Our anatomical and histochemical investigations indicate that the application of gibberellin can induce the formation of tension wood on vertical stems of angiosperm trees in the absence of gravitational stimulus.     

Modelling the distribution of diameter growth along the stem in Scots pine

This paper presents an empirical model for the distribution of diameter growth along the stem in Scots pine (Pinus sylvestris L.) and for the consequent stem form over time. First, the distribution of annual mass growth in the stem is determined as a function of the total annual growth in stem mass, current stem mass and the distribution of the latter along the stem. Second, the distribution of diameter growth is obtained by converting the fraction of annual growth in the stem mass at a given height in the stem into the thickness of the annual ring at the same height. Application of the model to Scots pine data sets including both young and mature trees not used in parameter estimation showed that the model was capable of reconstructing the distribution of diameter growth from the stem butt to the apex and from the pith to the stem surface at any height in the stem in both young and mature trees. The resulting empirical model was also linked to a physiological, process-based model in order to study its performance in a simulated stand. Simulations representing trees grown in unthinned and thinned Scots pine stands with trees of different status (from dominant to suppressed) showed that the response in tree growth to thinning in terms of the distribution of diameter growth along the stem was quite realistic relative to measured data.     

Daily dynamics in xylem cell radial growth of Scots pine (Pinus sylvestris L.)

Daily dynamics of radial cell expansion during wood formation within the stems of 25-year-old Scots pine trees (Pinus sylvestris L.), growing in field conditions, were studied. The samples of forming wood layers were extracted 4 times per day for 3 days. Possible variations in the growth on different sides of the stem, duration of cell development in radial cell expansion phase and dynamics of cell growth in this phase were taken into account. The perimeters of tracheid cross-sections as a reflection of primary cell wall growth were the criterion of growth in a radial direction. For the evaluation of growing cell perimeters a special system for digital processing and image analysis of tracheid cross-sections of the forming wood was used. Growth rate for certain time intervals was estimated by the change in the relation of the perimeter of each observed cell in each of ten tracheid rows in each of 12 trees to the perimeter of the xylem cell of the same row before the expansion. Temporal differences in average values of the relations were estimated by Analyses of Variance. The existence of daily dynamics of Scots pine xylem cell radial growth has been proved. Intensive growth of pine tracheids has been shown to occur at any time of the day and to depend on the temperature regime of the day and the night as well as water supply of stem tissues. Moreover, reliable differences (P = 0.95) in the increment of cell walls during tracheid radial expansion have been found. Pulsing changes of the water potentials both of the cell and the apoplast, as the reason for the fluctuations of radial cell growth rate, were discussed.     

Root deformation in plantations of container-grown Scots pine trees: effects on root growth,tree stability and stem straightness

Root system deformation was studied in 23 Scots pine (Pinus sylvestris L.) stands in central Sweden. The study comprised both plantations created with container-grown plants (Paperpot) and natural stands including young (7–9 year old) and older (19–24 year old) trees. Trees were measured with regards to distribution of roots, root deformation, stability, stem straightness and wood properties in stumps. Root distribution was most uniform for naturally regenerated trees. Older trees generally showed a better root distribution than young trees. The young planted trees displayed a high frequency of severely spiralled root systems, while only a few of the older trees had spiralled root systems. No severe root deformations were observed on naturally regenerated trees. Naturally regenerated trees were more stable than those which had been planted. Differences in bending moment, when trees were pulled to an angle of 10°, were considerable between young planted and naturally regenerated trees, but less pronounced for the older trees. Young planted trees had the highest frequency of severely crooked stem bases, while naturally regenerated trees had the straightest mode of growth. Tensile strength in peripheral wood samples of the stumps was substantially lower for planted than for naturally regenerated trees. Strain values to breakage of wood samples, taken from the root collar and the central- and peripheral part of the stump were lower for planted trees. The conclusions from this study are that root distribution, tree stability and stem straightness of planted Paperpot-grown trees will improve after a certain time and approach the state of naturally regenerated trees. As trees grow older, early established crooked stem bases will be compensated by radial growth and the tree will appear straighter. Inside the stem, however, problems may still remain with abnormal fibre direction and compression wood together with inferior root strength due to fibre disturbances as a result of spiralled roots. This revised version was published online in June 2006 with corrections to the Cover Date.