Cork oak (<Emphasis Type="Italic">Quercus suber</Emphasis> L.) wood growth and vessel characteristics variations in relation to climate and cork harvesting

Variations in tree ring growth of Quercus suber L. were analysed using dendrochronological techniques on cork oak discs from trees harvested in the cork producing region of Alentejo, Portugal. A tree-ring chronology containing a strong common signal and covering the period from 1970 to 1995 was build for ca. 30-year-old cork oaks never submitted to cork harvesting using 14 trees that crossdated satisfactorily out of 30 sampled trees. The tree ring indices correlated positively with September temperature (r = 0.48, P < 0.05) and very strongly with precipitation totals from previous October until current February (r = 0.82, P < 0.001) showing that the water stored in the soil during the autumn and winter months prior to the growing season has a primordial effect on the growth of the given season. The effects of cork harvesting were analysed by comparing mean ring width, mean annual vessel area, vessel density (n°vessels/mm²), and vessel coverage (percentage of transverse surface occupied by vessels) between three mature cork oak trees and three young trees, for the period from 1987 to 1996, corresponding to the growth between two consecutive cork removals in the case of mature trees. In 1988, 1989 and 1996 (corresponding to the first and second years after cork removal, and 1996 to a year of cork removal), the ratios between ring widths of young versus mature trees was twice that for the rest of the period. However, an effect of cork removal indicated by eventual alterations in vessel size and distribution in the wood rings corresponding to the years 1988, 1989 and 1996 in the mature cork oaks was not observed.     

Effects of radial growth rate on selected indices for juvenile and mature wood of the Japanese larch

To examine the differences between juvenile and mature wood, 12 aged sample trees from two areas of Nagano Prefecture were harvested; and the radial development of tracheid length, the ring density, and the relation of the radial growth rate (observed by ring width) with some selected indices of ring structure were investigated. The results proved that the radial variation of tracheid length with ring number can be described by a logarithmic formula, and both plantations reached the demarcation of juvenile and mature wood at age 18. With the segmented regression method, we also analyzed radial variation of mean density and found that the demarcation of juvenile and mature wood was at age 15 for sample trees from Saku and at age 21 for those from Yabuhara. By using the results of estimates from juvenile and mature wood based on ring density, we found that high growth rates resulted when producing lower-density wood during the juvenile period, but these rates did not occur during the mature period. The basic reason for this phenomenon is the variation in patterns of earlywood and latewood in juvenile and mature wood, respectively. This result advised us that when managing plantations of Japanese larch it is necessary to take different measurements at different growth periods.     

Xylem traits in European beech (Fagus sylvatica L.) display a large plasticity in response to canopy release

Key message

The position of trees in the canopy impacts xylem structure and its inter-annual variation. After canopy release, the increase in the hydraulic conductivity of growth rings was driven by an increase in radial growth in large trees, and by both an increase in radial growth and changes in xylem structure in saplings.

Context

Forest canopies are frequently subjected to disturbances that allow understory trees to access the upper canopy. The effect of canopy release on xylem anatomy has been assessed in juvenile trees and saplings, while the potential acclimation of larger trees remains poorly documented.

Aims

We estimated the potential hydraulic conductivity of growth rings in large understory trees compared to overstory trees, and evaluated the responses to canopy release in large trees and in saplings.

Methods

We recorded radial growth, wood density, and vessel structure in beech trees according to their position within the canopy and their size. Xylem traits were followed during 6 years after canopy release for large trees, and during 2 years for saplings. Vessel diameter and frequency as well as ring area were used to compute the potential annual ring hydraulic conductivity.

Results

Large understory trees displayed lower radial growth increments and lower potential annual ring hydraulic conductivity than overstory trees. After canopy release, potential annual ring hydraulic conductivity increased in large trees, due exclusively to increased radial growth without any change in specific hydraulic conductivity. It increased in saplings due to both increased radial growth and increased specific conductivity.

Conclusion

Tree size impacted xylem structure and resulted in plasticity of the potential hydraulic conductivity of the annual tree ring following canopy release.

     

Solar activity, global surface air temperature anomaly and pacific decadal oscillation recorded in urban tree rings

A tree ring width chronology developed from 58 samples taken from Chinese pine trees (Pinus tabulaeformis Carr.) growing in Shenyang city was studied to analyze the effects of solar activity, Global Surface Air Temperature Anomalies (GSATA) and the Pacific Decadal Oscillation (PDO) on annual radial growth. An excellent response of tree rings to solar activity and global environmental change was revealed, indicating that these urban Chinese pine trees are a suitable proxy for Sun-Earth system research despite their location in Shenyang, an industrial city. The information derived from annual growth rings of urban Chinese pine trees coincides with records from undisturbed trees in a natural location, and demonstrates significant synchronous response to solar activity in the periodic bands of 5–8, 10–12 and 20–30 years. The wave signal of tree growth was also affected and amplified by the combined effect of the PDO from 1900 to 2004 and GSATA from 1880 to 2004 with a short lag time, revealing 5–8, 11–13, and 20–23 year oscillatory modes. At 20–30 year timescale, Chinese pines validated the strong GSATA influence signal at 20.8 years, and may be the primary cause of the 20 and 23 year periodic waves of tree rings. Shenyang urban tree ring growth also showed high frequency variation (2—4 y) that may be due to low periodicity of solar activity, GSATA, PDO and local climate variations, especially in the 2–4 years band width.     

Radial variations of wood properties of an endangered species, <Emphasis Type="Italic">Pinus armandii</Emphasis> var. <Emphasis Type="Italic">amamiana</Emphasis>

A dead tree of Pinus armandii Franch. var. amamiana (Koidz.) Hatusima (abbreviated to PAAm) was obtained from a natural habitat on Tanega-shima Island and various properties of its wood were investigated. Grain angle was measured and soft X-ray analysis was undertaken to obtain the density in each annual ring. Unit shrinkage and dynamic properties were measured by shrinkage, bending, and compression tests. Variations of wood properties in the radial direction, relationships of wood properties to density, and annual ring width were examined. Roughly speaking, variations in the radial direction of the grain angle, twist angle by drying, Young’s modulus and strength in static bending, absorbed energy in impact bending, compressive Young’s modulus, compressive strength, and compressive proportional limit corresponded to the variation of annual ring width. As a result, it was determined that if PAAm is afforested artificially for the purposes of lumber production and conservation, the annual rings of logs should not be too widely spaced. Wood properties of PAAm were similar to those of Japanese black pine (Pinus thunbergii Parl.), which is another representative pine on Tanegashima Island. This study was presented in part at the 56th Annual Meeting of the Japan Wood Research Society, Hiroshima, August 2007     

Asynchronous wood formation in youngAcacia mangium planted in Malaysia

In general, tropical trees have less-distinct growth rings. Even if the trees have some concentric structures, there is little information on the timing of their formation. The objective of this report is to show the asynchronous wood formation from the observation of indistinct, ring-like structures. YoungAcacia mangium planted in Malaysia was investigated for asynchronous wood formation anatomically and macroscopically. Tree heights and girths at breast height for 28 trees were measured every other week. Wood disks from four trees were sampled at breast height for this research. The radial variation of vessel dimensions and the number of cambial cells in different radial directions and the wood anatomy of discontinuous bands were investigated using thin sectioning and soft radiographic methods. The vessel dimension was quite different in their radial variation even for four radial directions with a similar radius. There was a large variation in the number of cambial cells among radial directions in youngAcacia mangium. Bands of low density included a layer of vessels aligned in a spiral rather than in a closed circle.     

Effect of growth conditions on wood density of Spanish Pinus nigra

The natural forests of Laricio Pine (Pinus nigra Subsp. salzmannii) cover in Spain an area of 353,000 ha, producing annually 310,000 m³ of roundwood. This species is considered as the fourth, most important Spanish conifer, being its timber frequently used in construction. The natural area of distribution of Laricio pine in Spain is officially divided into ten different regions of provenance, being only four of them considered as productive. The present study on the influence of growth conditions on extracted wood basic density (EBD) is carried out on 89 P. nigra trees, sampled from the two most productive Spanish regions of provenance (R.7 and R.8) in seven different sites. Cross-sectional disks were cut at 3 m (in 89 trees) and every 3 m height (in 18 trees) from the butt up to 15 m. For assessing the pattern of radial variation of the properties, 20×20×20 mm³ wood samples were taken radially from every cross-sectional disk. In every 20×20×20 mm³ wood sample, age and distance to the pith of the central annual ring was recorded, together with the average growth rate. Extractive content and EBD were measured in every sample. The results lead to the conclusion that there is not a significant relationship between EBD and provenance or growth rate. On the contrary, a significant relationship has been found between EBD and site quality or cambial age. The results also demonstrate that in a tree EBD tends to diminish from the base to the top and from the pith outwards. The main result of the present study is related with the observation that for the same cambial age, annual ring width has no significant influence on EBD, confirming that its utility as a predictor of the quality of timber is highly questionable, at least if the ring curvature is not considered.     

Models for predicting microfibril angle variation in Scots pine

Context

Microfibril angle (MFA) is one of the key determinants of solid timber performance due to its strong influence on the stiffness, strength, shrinkage properties and dimensional stability of wood.

Aims

The aim of this study was to develop a model for predicting MFA variation in plantation-grown Scots pine (Pinus sylvestris L). A specific objective was to quantify the additional influence of growth rate on the radial variation in MFA.

Methods

Twenty-three trees were sampled from four mature Scots pine stands in Scotland, UK. Pith-to-bark MFA profiles were obtained on 69 radial samples using scanning X-ray diffractometry. A nonlinear mixed-effects model based on a modified Michaelis–Menten equation was developed using cambial age and annual ring width as explanatory variables.

Results

The largest source of variation in MFA (>90 %) was within trees, while between-tree variation represented just 7 % of the total. Microfibril angle decreased rapidly near the pith before reaching stable values in later annual rings. The effect of ring width on MFA was greater at higher cambial ages.

Conclusion

A large proportion of the variation in MFA was explained by the fixed effects of cambial age and annual ring width. The final model is intended for integration into growth, yield and wood quality simulation systems.     

Age and weather effects on between and within ring variations of number,width and coarseness of tracheids and radial growth of young Norway spruce

Annual growth, fibre and wood properties of Norway spruce are all under strong influence from genetics, age and weather. They change dynamically, particularly at young ages. Most genetic research and tree improvement programs are based on data from this most dynamic phase of the life of trees, affected by differences in weather among sites and years. In the work presented, influences of age and weather were investigated and modelled at the detail of annual rings and at the sub-tree ring level of earlywood, transitionwood and latewood. The data used were analysed from increment cores sampled at age 21 years from almost 6000 Norway spruce trees of known genetic origin, grown on two sites in southern Sweden. The traits under investigation were radial growth, cell widths, cell numbers, cell wall thickness and coarseness as a measure of biomass allocation at cell level. General additive mixed models (GAMMs) were fitted to model the influences of age, local temperature and precipitation. The best models were obtained for number of tracheids formed per year, ring width, average radial tracheid width in earlywood, and ring averages for tangential tracheid width and coarseness. Considering the many sources behind the huge variation, the explained part of the variability was high. For all traits, models were developed using both total tree age and cambial age (ring number) to express age. Comparisons indicate that the number of cell divisions and ring width are under stronger control of tree age, but the other traits under stronger control of cambial age. The models provide a basis to refine data prior to genetic evaluations by compensating for estimated differences between sites and years related to age and weather rather than genetics. Other expected applications are to predict performance of genotypes in relation to site or climate and simulation of climate change scenarios.     

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.     

Long-term growth analyses of Japanese cedar trees in a plantation: neighborhood competition and persistence of initial growth deviations

The individual growth of tree diameter at breast height (dbh) is analyzed in an even-aged plantation of Cryptomeria japonica from stand age of 45 to 94 years, to examine how the growth of individual trees has been affected by the changes in spacing resulting from thinning operations. At any age, a significant proportion (0.37–0.46) of the variation in dbh growth during a 5–11-year period was explained by dbh at the beginning of the period, probably due to greater leaf mass of larger trees. Next, either one-sided or two-sided competition was added to the model, by calculating the basal area (BA) of neighboring trees around each tree within a given radius or BA for trees having larger dbh than the focal tree within the radius. After preliminary analyses, a radius of 8 m was selected as the critical range for tree competition. Although both types of competition explained a significant proportion (0.09–0.43) of growth variation, one-sided competition was not significant at ages greater than 54 years. Based on the model at 45 years of age, the initial deviation of growth rate for each tree from the predicted rate was calculated and added to the models as a third variable. This raised the coefficient of determination up to 0.50–0.74. These findings have practical significance for forest plantation management, particularly for controlling the growth of standing trees via thinning, to produce high-quality timber in the future.     

The effect of bark decortication for hiwada production on xylem and phloem formation in Chamaecyparis obtusa

Of all plant materials used to cover the roofs of traditional Japanese buildings, Japanese cypress (Chamaecyparis obtusa) bark, hiwada, has the longest service life and has been used from ancient times. However, wood and bark properties after hiwada harvest have not been evaluated in detail. We studied whether decortication for hiwada production in winter affected xylem and phloem formation. Decorticated trees still preserved all inner bark and part of the outer bark, and both decorticated and control trees had similar annual ring structures at all stem heights in the xylem and phloem. In both xylem and inner bark, no significant difference in ring width at any stem height was found between annual rings before and after decortication. Thus, this study revealed that the decortication of bark for hiwada production does not affect the formation of xylem and the inner and outer bark if decortication is carried out by highly skilled workers in winter.     

Integration of pines and pastures for hay and grazing

When combining pine and cattle production on improved pastures, grazing may have to be delayed for several years until trees are large enough to resist injury. During this period forage would be lost unless harvested for hay. This study in south Georgia, USA, examined hay production during the first 3 years, cattle production during the next 3 years, and effects of this management system on survival and growth of slash pine(Pinus elliottii) planted in widely-spaced rows within the pastures. Pines were planted at 225 trees/ha in configurations of 3.0 x 14.6 or 4.9 x 9.1 m in pastures recently sprigged or seeded with Coastal bermudagrass (Cynodon dactylon) or Pensacola bahiagrass(Paspalum notatum). Pastures were fertilized annually and cut for hay four or five times each year. Wide-row spacings permitted haying operations, but tree rows and turning areas removed 5 to 8 percent of the area from production. However, hay yields and liveweight gains by yearling cattle were near normal for this locale. Slash pine planted in these fertilized pastures survived well, grew rapidly, and only a few trees were killed during haying operations. After 6 years, 83 percent of the trees survived and averaged 6.5 m in height and 13.2 cm in diameter. Cattle killed a few trees by rubbing during the 4th year. Wide-row spacing of pines permits harvesting forage for hay while waiting for the trees to become large enough to permit grazing. This approach to agroforestry produces annual returns to the landowner while awaiting maturity of the pines.     

Height growth of young larch (Larix kaempferi) in relation to the frequency of deer browsing damage in Hokkaido, Japan

The effect of browsing frequency by Sika deer (Cervus nippon yezoensis) on the height growth of Japanese larch (Larix kaempferi) in Hokkaido, Japan, was analyzed. Tree heights at the time of the census and at the start of the current growing season were determined for 100 trees selected randomly at each site. The tallest shoot on each tree was identified and the number of browsing occurrences on the current-year shoot was determined. Young larches browsed by deer repeatedly develop new shoots during a growing season. Larch is a browsing-tolerant species, and the average height growth of the most heavily browsed trees was >15 cm. Although resprouting partly compensated for decreased height growth caused by browsing, the annual increase in height decreased with increased browsing frequency. Thus, browsing frequency during a growing season best reflects the intensity of browsing damage to larch trees.     

Radial variation of wood density components and ring width in cork oak trees

The radial variation of ring width and wood density was studied in cork oaks (Quercus suber) using microdensitometry. The observations were made in young never debarked cork oaks (30–40 years of age) and in mature trees under cork production (37–60 years of age). The cork oak wood is very dense (mean ring density 0.86 g.cm^?3, between 0.79 g.cm^?3 and 0.97 g.cm^?3) with a small intra-ring variability (mean earlywood density 0.80 g.cm^?3 and latewood density 0.90 g.cm^?3). The density components decreased from pith to bark more rapidly until the 15th ring, and then only slightly. There were no significant differences in the mean density components between never debarked trees and trees under cork production but their outwards decrease was accentuated in the never debarked trees. The annual growth was high, with a ring width mean of 3.9 mm (4.2 mm in the first 30 years) and the latewood represented 57% of the annual growth.     

Variation in treatability of Scots pine sapwood: a survey of 25 different northern European locations

The treatability of Scots pine sapwood sampled from 25 locations in northern Europe, which was impregnated with an experimental furfuryl alcohol mix, was studied. A large variation in treatability was found between stands. The treatability was affected by anatomical properties of the trees and therefore also by the immediate climate, the sociological position of the tree in the stand, growth increments and on-stand competition. With the models applied, most parts of the differences in treatability between stands could be explained, while the treatability variation between trees within a stand remained mostly unexplained. Wider annual rings and higher latewood contents were found to increase the treatability. Models that included data on growth conditions and climate explain more of the variation in treatability, indicating that also other anatomical properties are influencing the treatability. Tree attributes and growth conditions that reduce annual ring width and latewood content decreased the treatability. Trees growing under warmer conditions and developing larger growth increments were easier to impregnate. Wood from trees growing near the timberline and under environmental conditions impairing wood growth was more difficult to treat.     

Effect of growth on wood properties for Japanese larch (Larix kaempferi): Differences of annual ring structure between corewood and outerwood

To investigate the relations between growth and the wood properties of Japanese larch (Larix kaempferi), six sample trees of varied ages and radial growth were felled and the ring width, ring density, percentage of latewood, and some other factors were determined. There were significant differences in ring density and percentage of latewood between sample trees with vigorous growth and those with poor growth. In corewood the ring density decreased with increasing ring width for all sample trees, whereas in outerwood this trend did not appear. Moreover, the latewood width increased with the increment of ring width only in outerwood, whereas there was almost no change in the corewood. The variation in patterns of ring width, ring density, and percentage of latewood in the radial direction and the relation with height was also studied.Part of this report was presented at the 47th annual meeting of the Japan Wood Research Society, Kochi, April 1997     

红豆树等6种珍贵用材树种的生长特性和材性分析

利用浙江龙泉和庆元两地在较好立地上营造的21 36年生红豆树、江南油杉、伯乐树、闽楠、刨花楠和乐东拟单性木兰6种珍贵用材树种的片林,以研究其生长、干形和木材基本密度的变化规律。研究结果表明,红豆树是一个树高和胸径生长量大、径生长速生持续期长、木材基本密度中等及其径向均匀性较高的珍贵用材树种,平均年轮宽度达到0.8 1.2 cm,36年生时按宽度和面积计算心材比例分别为60.57%和37.47%。因红豆树分叉干特性明显,在栽植的第2年就应及时修枝抹芽和施肥,以培育树干通直、心材比例高的优质干材。江南油杉和闽楠等其它5种珍贵用材树种皆较少分叉干,其树干通直。闽楠以材质优良而闻名,木材密度中等,虽然其生长速度较慢,但生长势很强,年生长量稳定,加之树冠窄小,适宜长周期大径材的培育;伯乐树生长速度中等,木材密度略低,但其径向均匀性较高,宜作为优质的工艺材来培育;树冠窄小的乐东拟单性木兰生长速度中等,但其木材密度大、径向均匀性高,是优良的珍贵用材树种;江南油杉是个早期生长快而后期生长慢,木材密度中等的针叶树种,可通过加强经营管理延长速生期和提高木材密度的径向均匀性;刨花楠虽然生长速度中等,但其木材密度却较低,浸水有粘液,不宜作为珍贵用材树种培育。     

Which factors most influence heartwood distribution and radial growth in plantation teak?

? To investigate the effect of climate on radial growth in young plantation grown teak (Tectona grandis L.), growth ring width was measured in 105 trees and correlated to precipitation and temperature data.

? The social status of trees within the stand was also determined and cross-sectional area (CSA) for the trunk correlated to the proportion of heartwood (HW) within the tree. HW develops asymmetrically in leaning stems of some conifer species, but it is not known if this phenomenon also occurs in broadleaf species. Therefore, we measured HW proportion in leaning and straight stems, along with the number of growth rings in the HW.

? Annual ring width depended strongly on mean monthly temperature during the rainy season and the most significant relationships were found corresponding to the months of June and July. With regard to the weaker relationship between precipitation and radial growth, correlations were highest during the period of bud-break at the beginning of the rainy season.

? The very high stand density affected radial growth, particularly in suppressed trees, which responded little to thinning operations. HW formation was greatest in dominant trees, and was highly regressed with stem CSA.

? Therefore, rapid growth of young stands should be encouraged by reducing stand density. Asymmetric HW formation occurred in both leaning and straight trees, and was significantly greater along the upper sides of leaning stems. It is probable that this eccentric HW formation is linked to mechanical loading on the tree.

     

Growth change of young Picea sitchensis in response to deer browsing

Taking advantage of the introduction of the black-tailed deer to the Queen Charlotte Islands (British Columbia, Canada), we used dendrochronological analyses to understand the consequences of deer browsing on Sitka spruce growth. We compared shape, radial growth, height growth and age of young spruce in three sites. We identified two types of trees growing side by side: (1) stunted and heavily browsed spruce, smaller than the browsing limit and (2) escaped spruce that were taller than the browsing limit but still browsed in their lower part. The compact and heavily ramified shape in stunted spruce was the result of repeated and intense browsing. In escaped spruce this was also the case below the browsing limit (1.16 m±0.07 m), in sharp contrast with the normal shape that escaped spruce resumed above the browsing limit. We show that the release of browsing pressure, once the tree reaches the browsing limit, is characterised by an abrupt increase in radial growth. Before release, trees show a growth stagnation characterized by narrow rings (0.5 mm per year) and small annual height increments (<5 cm per year). After release, trees show a growth stabilisation characterised by wider rings (3 mm per year) and larger annual height increments (20 cm per year). We use this pattern to estimate frequency and age at release and their possible variation over time. Age differences between stunted and escaped spruce are highly significant and indicate that, despite of browsing, most if not all trees will ultimately reach the browsing limit and escape. Heavy deer pressure (30 deer per km²) delays spruce sapling recruitment by about 8 years. This delay varies in relation to site quality and seems to have increased over time, suggesting an increase in browsing pressure.