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Trees DOI 10.1007/s00468-011-0675-2

ORIGINAL PAPER

Variations in the radial growth and wood density components in relation to cambial age in 30-year-old Pinus brutia Ten. at two test sites
Bilgin Guller Kani Isik Senay Cetinay

Received: 29 October 2011 / Revised: 19 December 2011 / Accepted: 23 December 2011 Springer-Verlag 2012

Abstract Radial growth and wood density are important traits in assessing wood quality. Our objective was to investigate patterns of variation of radial growth (ring width, earlywood width, latewood width, latewood proportion) and wood density (ring average density, earlywood density, latewood density) components in a 30-year-old Pinus brutia at two test sites in Turkey. Wood increment cores at a height of 1.30 cm (dbh) from 1,010 trees at age 30 years were evaluated at two test sites. The radial growth and wood density traits of the individual rings were measured using X-ray densitometry. The test sites showed statistically signicant differences in the radial growth traits but not in the wood density traits, suggesting that the wood density traits are less subject to environmental changes. The ring average density was relatively low (485 kg/m3) at early cambial ages (near the pith) and increased to 501 kg/m3 at later cambial ages (near the bark). The latewood density was 550 kg/m3 near the pith, increased steadily to 630 kg/m3 at cambial age 12, and remained stable thereafter. In contrast, the earlywood density and latewood proportion were highest near the pith. The twelfth ring from the pith appeared to represent the
Communicated by T. Fourcaud. B. Guller Department of Forest Products Engineering, Faculty of Forestry, Suleyman Demirel University, Isparta, Turkey K. Isik (&) Department of Biology, Faculty of Sciences, Akdeniz University, Antalya, Turkey e-mail: kani@akdeniz.edu.tr S. Cetinay Southwest Anatolia Forest Research Institute, Antalya, Turkey

transition from juvenile to mature wood. The unique relationships among early and latewood densities and latewood proportion in the juvenile and mature wood contribute to more uniform wood both within a given annual ring and between the juvenile and mature portions of the stem in P. brutia. Thinning increased the ring width, latewood proportion, and ring average density. Keywords Pinus brutia Ten. Cambial age Radial growth Wood density Juvenile wood Mature wood Thinning

Introduction Pinus brutia is naturally distributed in the eastern Mediterranean region, including Turkey, Greece, Cyprus, Syria, Israel, Palestine, Jordan, and Iraq (Boydak 2004). The species has been given high priority in plantation forestry in various countries with Mediterranean climates due to its relatively fast growth rate and wide ecological adaptability (Palmberg 1975; Fisher et al. 1986; Weinstein 1989). P. brutia has a wide geographic distribution in area (more than 5,500,000 ha forest land in eastern Mediterranean region) and altitudinal range, from sea level up to 1,400 m (asl), and is an important source of forest products in the region (Erkan 1998; Guller 2007). Radial and axial variations in wood density and its relationships with annual ring components (earlywood density, latewood density, and latewood percentage) are essential in assessing wood quality (Zobel and Sprague 1998; Savva et al. 2010). These traits are important for establishing the quality of pulp, paper, and sawn timber. They are also used to determine the transition age from juvenile to mature wood in forest m 2002). tree species (Tassisa and Burkhart 1997; Lindstro

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The portion of wood initiated near the pith at any height within a tree is called juvenile wood (JW). Mature wood (MW) is subsequently produced as the tree ages. Several physical properties (such as annual ring width, ring density, ber length, cell wall thickness, and micro-bril angle) and mechanical properties may differ between JW and MW (Zobel and Sprague 1998; Alteyrac et al. 2006). Wood density, a relatively easy trait to measure, is generally used to distinguish the JW-MW boundary. To our knowledge, information on wood density and annual ring components are limited for P. brutia. Adamopoulos et al. (2009) studied the ring width, latewood proportion, and dry density of 16 dominant P. brutia trees randomly chosen from two reforestation sites in Northeastern Greece. Two different methods for measuring the general wood density properties of P. brutia have been published (Guller 2010; Guller and Yasar 2010). However, there are no reports concerning the transition ages from juvenile wood to mature wood in the species. Moreover, additional information on radial growth and annual ring properties based on higher number of trees from the optimum distribution range of the species are needed. The purpose of this study was to investigate the radial variation of wood density and annual ring components in P. brutia. The specic objectives were to (i) investigate patterns of radial growth components (ring width, earlywood width, latewood width, and latewood proportion), (ii) determine the radial variation in wood density properties (ring average density, earlywood density, and latewood density) from pith to bark within trees, (iii) determine possible transition ages from juvenile to mature wood, (iv) examine the effects of test site on wood density and radial growth traits by comparing trees of the same age and similar seed sources planted at two test sites, and (v) examine the effects of thinning on the overall wood density and radial growth components.

Initially, except for the border trees, 1,800 trees were planted at each test site in 2.00 m 9 2.00 m distance in hexagonal arrangement. The understory vegetation was cleared manually at the test sites every 25 years (more often in early years) until crown closure. Two thinnings were applied at each test site prior to wood sampling [rst and second thinnings were performed at ages 13 (in Jan., 1991) and 17 (in Feb., 1995) at Dg, and at ages 17 (in Feb., 1995) and 28 (in May, 2007) at Kp, respectively]. After the second thinning, distances between trees were 3.46 m 9 3.46 m in hexagonal surroundings. The Duzlercami (Dg) site had a higher site index and growth rate than the Kepez (Kp) site (Table 1). Collection, care, and preparation of wood samples In August 2007, one increment core (12 mm thick) per tree was collected at breast height (1.3 m) in the northsouth direction from bark to bark, intersecting the pith. The trees were 30 years in age from seed (29 growing seasons in the eld) at the time of sampling. The diameter at breast height (dbh) for each tree was also measured. In total, 1,080 wood cores were collected from the two test sites with a minimum of six cores collected per family at each site. Immediately after removal, the increment core was stored in a cooler bag, subsequently vacuum-sealed in plastic bags, and stored at ?2C. For evaluation, the cores were divided into two radii and dried at room temperature. The radii of each core were glued to core holders (poplar strips) and 2-mm thick radial strips were cut. At the end of sample preparation process, a total of 1,024 clear samples were obtained. Variables (traits) studied The primary wood properties used in this study were components of annual ring width (radial growth) and annual ring density. These traits were abbreviated and dened as follows: WRW: whole annual ring width (width of an entire annual ring, mm); EWW: earlywood width (width of earlywood portion of an annual ring, mm); LWW: latewood width (width of latewood portion of an annual ring, mm); LWP: latewood proportion (proportion of an annual ring that is latewood, %); RAD: ring average density (mean density of a whole annual ring, kg/m3); RWD: ring area weighted density (calculated by weighting average ring density with ring area which was computed assuming a circular shape of a stem, kg/m3); EWD: earlywood density (density of earlywood portion of an annual ring, kg/m3);

Materials and methods Plant material and experimental sites The wood samples evaluated in this study were collected from 30-year-old trees planted in 1979 at two provenance test sites, Duzlercami (Dg) and Kepez (Kp) near Antalya City in southwestern Turkey (Table 1). The trees sampled at each test site represent six natural populations originating from different elevations ranging from 60 to 1,050 m asl in the Taurus Mountains along the Mediterranean Coast (Isik 1986; Isik et al. 2002). The experimental design at the test sites was single-tree plots in a randomized complete block with three interlocked replications, which allowed thinning as the trees aged (Libby and Cockerham 1980; Isik 1988).

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Trees Table 1 Description of the two test sites and sampled trees for wood study in Pinus brutia Attribute Test sites Kepez, Kpa Elevation (m, asl) Latitude Longitude Size of the test site Site indexb Mean rainfall (mm) (May; Aug.; Dec.; annual)c Total trees evaluated Number of rings evaluated per site per trait
a b c

Duzlercami, Dga 350 36580 2700 N 30320 5600 E 0.78 ha 19.8 30.7; 2.0; 261.3; 1,052.3 504 At least 9,168

90 36550 0000 N 30360 5400 E 0.78 ha 13.7 30.7; 2.0; 261.3; 1,052.3 506 At least 9,248

The subscripts g and p were adopted in the article to easily distinguish the relatively good (g) site Dg from the relatively poor (p) site, Kp Based on top height at age 25 (Usta 1991)

Climatic data from the nearest meteorological station [Antalya airport, 53 m asl; approximately 17 km (to Kp) and 25 km (to Dg), birds-eye view]

LWD: latewood density (density of latewood portion of an annual ring, kg/m3).

Measurement of radial growth and wood density components Annual ring widths and densities were determined using X-ray densitometry. A xed threshold density of 450 kg/m3 was used to set the boundary between earlywood and latewood within a ring (Guller 2010). The radial strips were conditioned to 8% equilibrium moisture content and scanned using an X-ray densitometer (Quintek Measurement Systems, Model QTRS-01X) integrated with a computer digital analysis system. The rst annual rings next to the bark of each sample were not evaluated because they were usually incomplete or damaged during bark removal. The X-ray attenuation, measured by the densitometer, was related to the density by ll = lm 9 q, where ll is the measured attenuation of the X-ray beam passed through the sample, lm is the sample mass attenuation coefcient, and q is the density. Therefore, the density calculation required knowledge of the mass attenuation coefcient (cm2/g) of the wood. The calibration to the appropriate mass attenuation coefcient was conducted using a set of 35 radial strips from cores with densities previously determined using the maximum moisture content method (Smith 1954). The 35 mass attenuation coefcients were averaged to provide the nal value used to calculate the wood density. Statistical analyses After excluding the cores with extreme readings, a total of 1,010 trees (1 core per tree) were used for the nal

statistical evaluation from the two test sites (Table 2). The number of rings for each core (each tree) ranged from 10 to 26 depending on the growth rate and age attained at breast height. Cambial age 1 (i.e., ring number 1) at the pith was not included in the analysis because of irregularities in the X-ray readings. Similarly, ring numbers 23, 24, 25, and 26 were not included in the nal evaluation due to relatively low sample sizes (B140) or uneven representation at the two test sites. Thus, the data used in the nal analysis included cambial age 2 (i.e., ring number 2, the closest ring to the pith, with sample size n2 = 948) through cambial age 22 (ring number 22, the closest ring to the bark, with sample size n22 = 289). The means and coefcient of variation (CV) for each trait were calculated for each test site and cambial age. The CV was a measure of variability for a given character. A character with a low CV value was less variable (more stable and uniform) across ring numbers and sites. The mean values and CVs of the traits were plotted against the ring numbers from pith to bark (referred to as cambial age proles). The ages of the transition from juvenile to mature wood were determined by visual interpretation of cambial age proles of ring density traits. Analysis of variance (ANOVA) was used to statistically compare test sites for all traits studied. For the ANOVA test of dbh, we used the following model: Yij l Si eij ; 1

where Yij is the dbh of the jth tree at the ith site; l is the overall mean; Si is the effect of ith site (i = 1, 2) with 2 variance r2 Si ; eij is the residuals with variance reij . For the ANOVA test of ring traits (whole ring width, earlywood width, latewood width, latewood proportion, ring average density, earlywood density, latewood

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Trees Table 2 The dbh, ring width and wood density statistics of Pinus brutia Traitsa Overall species Nb dbh (mm) WRW (mm) EWW (mm) LWW (mm) LWP (%) RAD (kg/m3) EWD (kg/m3) LWD (kg/m3)
a

Kepez site, Kp Mean SDc 161.1 60.70 2.93 1.63 1.58 0.95 1.38 0.93 46.93 14.57 495.45 44.99 388.14 24.15 619.78 55.40 Nb 506 9,523 9,248 9,530 9,422 9,632 9,439 9,611 Mean SDc 124.70 29.63 2.18 1.16 1.18 0.74 1.02 0.60 47.73 15.19 496.20 46.32 389.42 23.18 614.74 53.23

Duzlercami site, Dg Nb 504 9,310 9,175 9,302 9,168 9,327 9,194 9,313 Mean SDc 197.64 61.89 3.70 1.68 1.99 0.97 1.74 1.06 46.10 13.86 494.69 43.55 386.82 25.05 624.99 57.10

1,010 18,833 18,423 18,832 18,590 18,959 18,633 18,924

dbh Diameter at breast height, WRW whole ring width, EWW earlywood width, LWW latewood width, LWP latewood proportion, RAD ring average density, EWD earlywood density, LWD latewood density Number of observations SD standard deviation

b c

density), we compared test sites and cambial ages with similar ring numbers [for example, a given trait of cambial age 2 (ring no 2) at site Kp was compared with the corresponding cambial age trait at site Dg]. The model was as follows: Yijk l Si Rji eijk ; 2

Results Overall characterization in Pinus brutia Overall, the mean diameter at breast height (dbh) of 30-year-old P. brutia trees from the two test sites was 161.1 mm (Table 2). The overall mean for the whole ring width (WRW) was 2.93 mm, with relatively high variability (CV = 55%). The overall mean for early wood width (EWW) was 1.58 mm and for latewood width (LWW) was 1.38 mm. Latewood proportion (LWP) averaged 46.9% and was less variable (CV = 31%) than dbh and ring width characters. The overall mean for the ring average density (RAD) of breast height samples from the two test sites was 495.45 kg/m3. The mean latewood density (LWD) of 619.78 kg/m3 was approximately 60% larger than the mean earlywood density (EWD) of 388.14 kg/m3. The ring density components (EWD, LWD, and RAD) had markedly lower variability than the ring width components (EWW, LWW, and WRW) (Fig. 1). The variability of LWP was in between these two groups. Because of the strong relationship observed between the RAD and the ring area weighted density (RWD) on an individual core basis (r = 0.92, p \ 0.0001, n = 1,010), we only used the ring average density (RAD) trait in our subsequent analyses. Relation of ring width (radial growth) and its components to cambial age The WRW was highest at cambial ages 2 and 3 (3.8 mm at Kp, and 5.3 mm at Dg) (Fig. 2a). The ring width (radial growth) subsequently declined gradually until cambial age 8

where Yijk is the kth observation in the jth ring (jth cambial age) at the ith site; l is the overall mean; Si is the effect of ith site (i = 1, 2); Rj(i) is the effect of jth cambial age within the ith site (j = 121, i.e., ring numbers 2 through 22); and eijk is the residuals. We assumed that the site effects were xed and the cambial ages (ring numbers) were nested within sites. The 2 variance components [r2 Si , variances due to sites; rRji due to cambial ages within sites, r2 eijk due to residuals] for each ring trait were estimated using the restricted maximum likelihood (REML) method of PROC VARCOMP (SAS Institute 1990). We also performed ANOVA tests to compare the overall (sites pooled) and individual juvenile and mature wood within each test site. In addition, we compared the juvenile wood (JW) in Kp with JW in Dg and mature wood (MW) in Kp with MW at Dg. In all the three comparisons, the model was as follows: Yij l Mi eij ; 3

where Yij is the value of the jth ring at the ith maturity level; l is the overall mean (or, l is the site mean in the case of within-site comparisons); Mi is the effect of ith maturity level (i.e., juvenile or mature, i = 1, 2) (or, for the third comparison JW at Kp vs. JW at Dg, and MW at Kp vs. MW at Dg).

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Dg Kp

6
Dg-WRW Dg-EWW

60 50
5

Kp-WRW Kp-EWW Kp-LWW

Dg-LWW

CV (%)

40
4

30 20 10

Width mm
RA D D LW D LW W LW P EW W W R EW W

0
1

Fig. 1 Variability prole to compare variability levels for ring width and density traits in Pinus brutia at two different test sites (Kp Kepez, Dg Duzlercami)

0 0 2 4 6 8 10 12 14 16 18 20 22 24

Ring no (pith to bark)

b
and remained approximately at 2.0 mm at Kp and at 3.5 mm at Dg. At cambial age 14, the radial growth began to gradually decline again to a mean value of 1.4 mm at Kp and 2.7 mm at Dg until cambial age 22. Although the magnitudes were different between the poor (Kp) and good (Dg) sites, the trends within the cambial ages were similar at both sites. The EWW and LWW showed parallel trends to that for the WRW. Except for the rst two rings at Dg site, the EWW for all cambial ages were consistently larger than (or rarely equal to) the LWW at both test sites (Fig. 2a). The general trends in ring width components were similar at the two test sites, and the good site (Dg) consistently had larger values at each ring than the poor site. The variability (i.e., CV) for WRW was initially low (30% at Kp, 35% at Dg), but starting at cambial age 4, gradually increased to 47% at cambial age 20 (Fig. 2b) with regular oscillations. The WRW had lower variability than its components (EWW and LWW) at both test sites. The variability of EWW at any given cambial age at the poor site (Kp) was higher than that at the good site. The rate of change in the variability from early to later cambial ages was greater in the WRW than in each of its components (Fig. 2b). Relation of ring density and its components to cambial age The RAD was comparatively low (approximately 485 kg/m3) at early cambial ages, reaching a mean value of 501 kg/m3 at cambial age 22 (Fig. 3a). The RAD values gradually increased from pith to bark and were more or less similar at both test sites at any given cambial age. The EWD was greatest (approximately 410 kg/m3) at the early cambial ages, declining steadily during the next decade to a low of 380 kg/m3 (Fig. 3a). Thereafter, the EWD became more or less stable, reaching 383 kg/m3 at

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Dg-WRW Dg-EWW Kp-WRW Kp-EWW

60

50

CV (%)
40 30 20 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24

Ring no (pith to bark)

Fig. 2 a Cambial age prole for mean values of ring width traits in Pinus brutia at two different test sites (Kp Kepez, Dg Duzlercami). b Cambial age prole for the coefcient of variation (CV, %) values of ring width traits in Pinus brutia at two different test sites

cambium age 22. The general trend of EWD variation from pith to bark was parallel, and the values were similar at both test sites at any given cambial age. The LWD was lowest (approximately 550 kg/m3) at early cambial ages, increasing sharply during the next decade at both test sites (Fig. 3a). By cambial age 12, both sites had nearly similar values and trends. However, after cambial age 12, the LWD became nearly stable with a mean of 628 kg/ m3 at the poor site (Kp) and a mean of 660 kg/m3 at the good site (Dg). At early cambial ages, the LWD was greater at the poor site than at the good site. However, starting at cambial age 11, the LWD at the good site was consistently higher than at the poor site. The differences between the poor and good sites increased with increasing age. The RAD variability (CV) gradually increased from younger to older cambial ages at both test sites (Fig. 3b).

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a 700
Dg-RAD Dg-EWD Dg-LWD Kp-RAD Kp-EWD Kp-LWD

56 54 52 Dg-LWP Kp-LWP

650

600

Density kg/m 3

550

LWP (%)
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23

50 48 46

500

450

44 42 40 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23

400

350

Ring no (pith to bark)

Ring no (pith to bark)

b
12
Dg-RAD Kp-RAD Kp-EWD Kp-LWD

b 38
Dg-EWD Dg-LWD

Dg-LWP Kp-LWP

11 10 9

36 34 32

8 7 6 5 4 3 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23

CV (%)
Ring no (Pith to bark)

CV (%)

30 28 26 24 22 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22

Ring no (pith to bark)

Fig. 3 a Cambial age prole for mean values of ring density traits in Pinus brutia at two different test sites (vertical dashed line indicates the transition age from juvenile wood to mature wood). b Cambial age prole for the coefcient of variation (CV, %) values of ring density traits in Pinus brutia at two different test sites

Fig. 4 a Cambial age prole for the mean value (%) of latewood proportion (LWP) in Pinus brutia at two different test sites. b Cambial age prole for the coefcient of variation (CV, %) of latewood proportion (LWP) in Pinus brutia at two different test sites

After cambial age 12, the RAD variability was less at the good site than at the poor site. The EWD had the lowest variability among all the wood traits studied. As with other density traits, the CV was lower at early cambial ages and increased, but with a less steep trend, from 4.5% (at ring number 2) to 6.3% (at ring number 22) (Fig. 3b). The wood density variation for LWD increased at both test sites from younger to older cambial ages (Fig. 3b). The LWD was consistently less variable (more uniform) at the good site than at the poor site. The late wood proportion (LWP) was highest (approximately 54%) at early cambial ages, declining gradually to a mean of 46% at cambial age 22 (Fig. 4a). Thus, the LWP showed a declining trend from pith to bark as the trees aged. At earlier cambial ages (until the 6th ring from the pith), the LWP in a given annual ring was consistently higher at the good site than the poor site. However, starting

at cambial age 7, the LWP at the poor site was consistently higher than at the good site. Thus, the proportion of wood formed early in a growing season within an annual ring was higher at the good site than at the poor site. After cambial age 7, the general trend of LWP variation from younger cambial ages to older cambial ages was nearly parallel at both test sites. The variability of the LWP proportion was high at early cambial ages (3237%) and gradually declined after ring number 10 (Fig. 4b). The LWP was consistently less variable at the good site than at the poor site, which was the case also for LWD trait. Comparisons of test sites in terms of radial growth and wood density traits The test sites differed signicantly in radial growth traits (dbh, WRW, EWW, LWW) (Table 3). The values of radial

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growth traits were clearly larger at the good site (Dg) compared with those at the poor site (Kp) (Table 2). For example, the partitioning of variance components indicated that 53% of the variation in dbh was due to site differences. Similarly, 33% of the variation in WRW, 28% in EWW, and 23% in LWW were also due to site differences. For each of these characters, the differences due to cambial ages (within sites) had smaller variance components than those of the site differences, the corresponding values being 20, 12, and 22%, for WRW, EWW, and LWW, respectively (Table 3). The late wood proportion (LWP) differences between the sites were not statistically signicant. As a result, the variance in LWP due to site differences was less than 1% (Table 3). Similarly, the contribution of variance due to cambial age differences in LWP was low (3.6%). The wood density traits (RAD, EWD, LWD) at the two sites were nearly equal with no statistically signicant differences (Tables 2, 3). Less than 1.11% of variance in each of the RAD, EWD, and LWD traits was accounted for by site differences. However, the cambial ages within the sites were signicantly different, as illustrated by the cambial age proles for each trait shown in Fig. 3a. The contribution of variance due to cambial ages in RAD was the smallest (3.8%) among the wood density traits (Table 3). Juvenile wood (JW)mature wood (MW) transition Several features of wood density and radial growth traits in our study indicate that the cambial ages (i.e., ring numbers) between 11 and 13 years might be considered the transition zone between juvenile and mature wood in P. brutia. For example, the LWD increased steadily from ring number 2 until ring number 12, after which it was stable at 646 kg/m3 (Fig. 3a). Similarly, the EWD and RAD values were nearly stable following ring number 13 at 384 and 505 kg/m3, respectively (Fig. 3a). The CV for EWD was also stable after ring number 12 (Fig. 3b); the EWD was more uniform among the rings after cambial age 12. The LWP declined steadily until cambial age 912 and then gradually increased, remaining at 4648% (Fig. 4a). The mean values for each of the ring width components were relatively high at ring 2, declined steadily until ring numbers 810, and were more or less stable after ring number 12 (Fig. 2a). Although site Kp (lower site quality) appeared to have an earlier rate of change from the juvenile to mature wood compared with site Dg, the general trends were nearly similar at both test sites. These observations suggest that the transition from JW to MW in P. brutia occurs at the 12th year from the pith. Using the 12th cambial age as the transition age between juvenile and mature wood, we compared the JW

and MW portions of the core samples at each test site for ring width and density traits (Table 4; Fig. 5). The radial growth traits (EWW, LWW and WRW) were signicantly higher in the JW than in the MW (Fig. 5a) overall and at both test sites. Contrary to the ring width traits, two of the ring density traits (RAD, and LWD) were signicantly higher in the MW than in the JW overall and at both test sites. The EWD, however, differed from the two other ring density traits because it was higher in the JW than in the MW. The LWP was similar to the EWD with the exception that the differences between the LWP in the JW and MW were not signicantly different at the Kp test site (Table 4). Effects of thinning on radial growth and wood density Thinning appeared to stimulate certain radial growth and density traits of trees during the subsequent growing seasons. For example, at test site Dg, the LWW increased from 1.3 to 1.8 mm 1 year after the second thinning (in the 1996 growing season) with relatively high values in the subsequent years (Fig. 6). The EWW (from 1.9 to 2.4 mm) and WRW (from 3.2 to 4.2 mm) exhibited similar responses. The response to thinning at the good site (Dg) was more pronounced than that at the poor site (Kp). The LWP and RAD also increased markedly at the Dg test site in the years following thinning.

Discussion Overall wood density and radial growth values for Pinus brutia Using X-ray densitometry, our results showed that the overall wood density (i.e., RAD) in 30-year-old P. brutia trees was 495 45 kg/m3. This value was consistent with previous ndings measured using other methods (Guller 2007; Guller and Yasar 2010). The mean LWD (620 24 kg/m3) was 60% higher than the mean EWD. This difference might be due to various anatomical modications and variations in the structure and chemical composition between the cell walls of late and earlywood (Decoux et al. 2004). In particular, the high resin content in P. brutia wood could be an important contributing factor to higher latewood density (Raymond et al. 2004). The radial growth (as expressed by WRW, EWW, LWW) was higher in the annual rings closer to the pith than those closer to the bark. The radial growth declined sharply until cambial age 8 and subsequently remained stable. Studies by Adamopoulos et al. (2009) on P. brutia also indicate that the annual rings in early years are broader than those formed in later years.

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Trees Table 3 Analyses of variance (ANOVA) results comparing the two test sites and cambial ages within test sites for growth and wood density traits in Pinus brutia Traitsa dbh (mm) WRW (mm) Source of variation Between sites Within Between sites Cambial ages (site) Within EWW (mm) Between sites Cambial ages (site) Within LWW (mm) Between sites Cambial ages (site) Within LWPc (%) Between sites Cambial ages (site) Within RAD (kg/m3) Between sites Cambial ages (site) Within EWD (kg/m3) Between sites Cambial ages (site) Within LWD (kg/m3) Between sites Cambial ages (site) Within df Degrees of freedom
a b

dfb 1 1,009 1 40 18,791 1 40 18,381 1 40 18,790 1 40 18,548 1 40 18,917 1 40 18,591 1 40 18,882

Mean square 1,343,402.7 2,351.2 8,912.67 271.22 1.49 2,469.00 51.48 0.631 1,972.86 95.52 0.539 1.938 0.576 0.031 2,654.63 37,468.70 1,948.25 24,942.86 33,821.20 510.24 724,871.59 509,383.88 1,970.58

Prob. \0.0001 \0.0001 \0.0001 \0.0001 \0.0001 \0.0001 \0.0001 0.0741 \0.0001 0.7915 \0.0001 0.3956 \0.0001 0.2399 \0.0001

% variance component 53.04 46.96 32.55 20.27 47.18 28.05 12.28 59.67 23.01 22.09 54.90 0.53 3.61 95.86 0.00 3.83 96.17 0.00 12.36 87.64 1.11 35.27 63.62

See Table 2 for abbreviations For LWP, ANOVA test was applied using arsin transformation

The CVs for the density traits (ranging from 4.6 to 10.1%) were markedly smaller than those of the radial growth traits (ranging from 28.0 to 65%). Thus, the wood density components were less variable (more uniform) than the radial growth traits. This result suggests that in operational forestry, although there might be greater variation among trees in terms of their growth rates, the trees exhibit less variation in terms of their wood density traits. Wood density versus cambial age The RAD in P. brutia increased with increasing cambial age. This is consistent with previous observations in many conifers (Zobel and Sprague 1998). Decoux et al. (2004) suggested that this observation might be due to an increase in the thickness of the cell wall and the number of cell rows in the radial and tangential directions as the tree become older. In our study, the LWD (starting at ring number 2) and LWP (starting at ring number 9) also increased from pith to bark. However, the EWD was essentially constant from the 9th ring to the bark. Thus, the increase in RAD

from pith to bark appears to be primarily related to LWD and LWP. Jyske et al. (2008) reported an increasing wood density from the pith outward that was also related to increasing latewood density and proportion in Norway spruce. Effects of site differences The trees at the good site (Dg) had 60% more radial growth than that at the poor site (Kp). In addition, the LWD, RAD and LWP were generally more uniform at the good site than at the poor site. This difference was more pronounced in the mature wood portion. These results suggest that good sites produce not only higher quantity but also higher quality (more uniform) wood than poor sites. Adamopoulos et al. (2009) reported that the ring width and dry wood density traits from P. brutia samples taken from the base of trees were signicantly higher at a good site than at a moderate site, although the differences between the sites were small. In contrast, we did not nd any signicant differences between sites in wood density

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Trees
44.62 B 12.06 (4,272) 504.26 B 43.25 (4,352) 380.69 B 24.20 (4,289) 660.08 B 45.27 (4,337)

Table 4 Means (standard deviations) of the radial growth and wood density traits in juvenile (JW, rings numbers 2 through 11) and mature wood (MW, rings numbers 12 through 22) in Pinus brutia

For a given trait, the JW and MW means that have different uppercase letters within the same test site (or within the overall species level) are signicantly different at least at the 5% level

Data are based on individual ring values. N number of rings (number of observations) are given in parenthesis

382.96 B 24.23 (8,780)

46.22 B 13.75 (8,751)

503.24 B 46.54 (8,961)

644.90 B 51.46 (8,930)

2.39 B 1.39 (8,976)

1.34 B 0.85 (8,756)

1.08 B 0.68 (8,960)

traits (RAD, EWD, LWD) despite large differences between the sites in radial growth. The least amount of variance explained by site factors was 23.0% for the radial growth components (WRW, EWW, LWW), while the corresponding values for the wood density components (RAD, EWD, LWD) and LWP were much smaller, less than 1.2%. Our results indicate that the higher annual growth increment associated with better site conditions does not necessarily result in a lower wood density. Nyakuengama et al. (2002) reported that for P. radiata, the ring width is more responsive than the wood density to fertilizer application. Berges et al. (2008) also reported that the variances due to site factors were twice as high for the radial growth components as for the wood density components in Quercus petraea from locations with different site indices. The results of these and several other studies (Zobel and van Buijtenen 1989) support our ndings on P. brutia that environmental factors affect the radial growth characters more than the wood density characters. The variances due to sites were zero or near zero for the EWD and RAD traits and was only 1.1% for LWD and not signicant. These results indicate that the LWD is more sensitive to environmental differences than the EWD and RAD. Indeed, although both test sites were similar in terms of the EWD and RAD up to cambial age 12, the LWD values at Kp were smaller than those at Dg at ages greater than 12 (Fig. 3a). Furthermore, as observed in Fig. 3b, the LWD was more variable at Kp than Dg, and trends from pith to bark for LWD were steeper (i.e., more sensitive) than other density traits. Raiskila et al. (2006) also observed that the latewood components are relatively sensitive to environmental and site effects within clones of Picea abies. Juvenile versus mature wood In this study, several features of wood density and radial growth traits indicated that the transition period between juvenile and mature wood in P. brutia occurred at cambial age 12. At cambial age 12 (ring number 12), density traits became nearly stable. This period started approximately 2 years earlier at the poor site (Kp) compared with the good site (Dg). In Pinus taeda, cambial ages 812 were considered to be the transition period between juvenile and mature wood where the wood that was produced after cambial age 11 exhibited considerably different characteristics than the wood produced at earlier ages (Megraw 1985). Burdon et al. (2004) suggested that rings 1115 from the pith represented the transition period for Pinus radiata. The general consensus in conifers has been that the quality of the juvenile wood is much poorer than the mature wood (Zobel and Sprague 1998; Alteyrac et al.

3.19 B 1.46 (4,345)

1.81 B 0.89 (4,275)

MWd

1.42 B 0.75 (4,340)

47.39 A 15.14 (4,896)

486.32 A 42.07 (4,975) 502.28 B 49.43 (4,609) 490.61 A 42.52 (5,023)

392.18 A 24.55 (4,905) 385.13 B 24.07 (4,491) 393.32 A 21.61 (4,948)

47.75 A 15.04 (4,479)

47.71 A 15.33 (4,943)

600.26 A 49.33 (5,018)

2.69 A 1.24 (4,892)

1.44 A 0.82 (4,767)

Kepez site, Kpc

JWd

1.26 A 0.66 (4,910)

630.57 B 52.83 (4,593)

1.65 B 0.76 (4,631)

0.90 B 0.50 (4,481)

MWd

0.77 B 0.39 (4,620)

594.40 A 48.03 (4,976)

4.15 A 1.73 (4,965)

2.14 A 1.01 (4,900)

Duzlercami site, Dgc

JWd

2.03 A 1.21 (4,962)

EWD (kg/m3) (N)

LWD (kg/m3) (N)

RAD (kg/m3) (N)

WRW (mm) (N)

EWW (mm) (N)

LWW (mm) (N)

LWPb (%) (N)

Traitsa

See Table 2 for abbreviations

For LWP, the ANOVA test was applied using arsin transformation

Overall speciesc (sites pooled)

MWd

47.55 A 15.24 (9,839)

488.48 A 42.35 (9,998)

392.75 A 23.13 (9,853)

JWd

597.34 A 48.77 (9,994)

3.42 A 1.67 (9,857)

1.80 A 0.99 (9,667)

1.64 A 1.05 (9,872)

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Trees

J
4

M
3

Width mm

J J

J M J M M J M M

0 Dg Dg Kp Kp Dg Dg Kp Kp Dg Dg Kp Kp

EWW

LWW
M M J J

WRW

b 680
660 640 620 600 580 560 540 520 500 480 460 440 420 400 380 360 340 320 300 Dg Dg Kp Kp Dg

kg/m3

M J J

Dg

Kp

Kp

Dg

Dg

Kp

Kp

EWD

LWD

RAD

Fig. 5 a Chart to compare the ring width traits (EWD, LWW and WRW) in juvenile (J) and mature wood (M) in Pinus brutia at two different test sites. b Chart to compare the wood density traits (EWD,

LWW and WRW) of juvenile (J) with mature wood (M) in Pinus brutia at two different test sites (Dg Duzlercami, Kp Kepez)

2006). In P. brutia, we found that the LW formed in juvenile wood was signicantly lighter than the LW formed in mature wood, as might be expected. In contrast, the EW formed in juvenile wood was signicantly denser

than the EW formed in mature wood. Furthermore, the LWP in juvenile wood was also signicantly higher than the LWP in mature wood. These relationships among EWD, LWD, and LWP in the juvenile and mature wood

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Trees
7 Dg-WRW 6 Kp-WRW Dg-EWW Kp-EWW Dg-LWW Kp-LWW

Only Dg
5

Both Dg and Kp

Width mm

8 19

19

85

8 19

8 19

8 19

19

89

90 991 992 993 994 995 996 997 998 999 000 001 002 003 004 005 006 007 2 2 1 1 1 1 1 1 1 19 2 2 2 2 2 2 1 1

Year

Fig. 6 Trends in latewood (LWW), earlywood (EWW) and ring (WRW) widths over the years in Pinus brutia at two different test sites (Kp Kepez, Dg Duzlercami). Arrows and vertical dashed lines indicate thinning years

are unique to P. brutia and contribute to a more uniform wood (and thus, improved wood quality) within a given annual ring and between the juvenile and mature wood portions of the stem. As a result, the difference between the RAD values in the mature and juvenile wood in P. brutia was not large. Effects of thinning

Conclusions The ring growth components (whole ring growth, earlywood growth, and latewood growth) were higher in the rings closer to pith than those near the bark. Thinning appeared to increase ring width, latewood proportion, and ring average density in subsequent growing seasons. The ring density traits (ring average density, earlywood density, latewood density) were much less variable (and less subject to changes by environmental variables) than the ring growth traits. The ring average density showed a gradual increase from pith to bark. This increase appears to be primarily related to latewood density and latewood proportion. Test site differences did not inuence the magnitude of the ring density traits signicantly. However, the ring density traits were more uniform among the rings at the good site compared with the poor site. An increased radial growth associated with better site conditions does not necessarily have an adverse effect on the wood density. Improving site quality would result in more uniform wood. Several features of wood density and radial growth traits reveal that the transition between juvenile and mature wood in P. brutia might occur at cambial age 12. This transition process takes place a few years earlier at poor sites compared with good sites.

We found no clear effect of the rst thinning on tree growth at either test site. This observation was probably due to the competition among trees had not yet started at the time of the rst thinning. By the time of the second thinning, however, the crown closure and competition among the trees were well advanced at the good site, Dg. As a result, the response to thinning at the good site was more pronounced than that at the poor site, Kp. The ring width and latewood proportion increased during the four subsequent growing seasons after the second thinning at Dg. Studies of P. brutia plantations indicate that thinning, especially heavy thinning, signicantly increased the radial growth rate (Guller 2007). Similarly, Jaakkola et al. (2006) showed that thinning in Picea abies stands signicantly increased the radial growth rate of the individual trees but had no or only a slight effect on the wood density. The ring average density (RAD) in the years following the second thinning at site Dg also increased, which appears to be an indirect effect of the increase in the latewood proportion.

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Trees

The radial growth traits (WRW, EWW, and LWW), LWP and EWD had signicantly greater values in the JW than in the MW. In contrast, two ring density traits, RAD and LWD, were signicantly higher in the MW than in the JW. Such unique relationships among LWP, EWD, and LWD in juvenile and mature wood contribute to more uniform wood (i.e., better quality wood) both within a given annual ring and between the juvenile and mature wood portions of the stem in P. brutia.

Acknowledgments The study was supported by The Scientic and Technological Research Council of Turkey (TUBITAK) under Project No: 106O442 and also by a post-doctoral research grant to Dr. Bilgin Guller at North Carolina State University (NCSU), Raleigh, NC, USA. We thank Dr. Steve McKeand, Dr. Fikret Isik and the staff of the NCSU Tree Improvement Program for their helpful suggestions and for providing X-ray facilities. Yusuf Kurt, Adnan Guller and Asli Gocmen helped during the sampling of the wood cores, and Dr. Brad St. Clair (USDA Forest Service, Corvallis, Oregon) thoroughly revised the language of the manuscript. Dr. Kani Isik established the test sites in 1979 and since then, the staff of the Forest Service and the Southwest Anatolia Forest Research Institute in Antalya has maintained the test sites. The Akdeniz University Scientic Research Fund and Suleyman Demirel University provided partial support and laboratory facilities. Two anonymous reviewers made precious suggestions on an earlier draft of the manuscript. The authors are grateful to all these persons and institutions.

References
Adamopoulos S, Milios E, Doganos D, Bistinas I (2009) Ring width, latewood proportion and dry density in stems of Pinus brutia Ten. Eur J Wood Prod 67:471477 Alteyrac J, Cloutier A, Zhang SY (2006) Characterization of juvenile wood to mature wood transition age in black spruce (Picea mariana (Mill.) BSP) at different stand densities and sampling heights. Wood Sci Technol 40:124138 Berges L, Nepveu G, Franc A (2008) Effects of ecological factors on radial growth and wood density components of sessile oak (Quearcus petraea Lielbl.) in Northern France. For Ecol Manag 255:567579 Boydak M (2004) Silvicultural characteristics and natural regeneration of Pinus brutia Ten.a review. Plant Ecol 171:153163 Burdon RD, Kibblewhite RP, Walker JCF, Megraw RA, Evans R, Cown DJ (2004) Juvenile versus mature wood: a new concept, orthogonal to corewood versus outer wood with special reference to Pinus radiata and P. taeda. For Sci 50(4):399415 Decoux V, Varcin E, Leban JM (2004) Relationships between the intra-ring wood density assessed by X-ray densitometry and optical anatomical measurements in conifers: consequences for the cell wall apparent density determination. Ann For Sci 61:251262 Erkan N (1998) Fast growing species and Pinus brutia. Workshop, Orman Bakanligi Yayin Dairesi Baskanligi no:63, Ankara (in Turkish, English abstract) Fisher JT, Neuman RW, Mexal JG (1986) Performance of Pinus halepensis/brutia group in southern New Mexico. For Ecol Manag 16:403410

Guller B (2007) The effects of thinning treatments on density, MOE, MOR and maximum crushing strength of Pinus brutia Ten. wood. Ann For Sci 64:467475 Guller B (2010) Determining wood density of Turkish red pine (Pinus brutia Ten.) by using X-ray densitometer. SDU For Fac J 2:97109 Guller B, Yasar S (2010) Estimation of Pinus brutia Ten. wood density from Fourier Transform Infrared (FTIR) spectroscopic bands by Articial Neural Network (ANN). Sci Res Essays 5:17651769 Isik K (1986) Altitudinal variation in Pinus brutia Ten.: seed and seedling characteristics. Silvae Genet 35:5866 Isik K (1988) Interlocked honeycomb design with three replications and its application on a population genetics analysis of Turkish red pine (Pinus brutia Ten.) (in Turkish, English summary). Turk J Agric For 12:377386 Isik F, Isik K, Yildirim T, Li B (2002) Annual shoot growth components related to growth of Pinus brutia. Tree Physiol 22:5158 kinen H, Saranpa a P (2006) Wood density of Norway Jaakkola T, Ma spruce: responses to timing and intensity of rst commercial thinning and fertilization. For Ecol Manag 237:513521 kinen H, Saranpa a P (2008) Wood density within Norway Jyske T, Ma spruce stems. Silva Fenn 42:439455 Libby WJ, Cockerham CC (1980) Random non-contiguous plots in interlocking eld layouts. Silvae Genet 29:183190 m H (2002) Intra-tree models of juvenile wood in Norway Lindstro spruce as an input to simulation software. Silva Fenn 36:521534 Megraw RA (1985) Wood quality factors in loblolly pine. Tappi Press, Atlanta Nyakuengama JG, Downes GM, Ng J (2002) Growth and wood density responses to later-age fertilizer application in Pinus radiata. IAWA J 23:431448 Palmberg C (1975) Geographic variation and early growth in southeastern semi-arid Australia of Pinus halepensis Mill. and Pinus brutia species complex. Silvae Genet 24:150160 a P, Fagerstedt K, Laakso T, Lo ija M, Mahlberg Raiskila S, Saranpa R, Paajanen L, Ritschkoff AC (2006) Growth rate and wood properties of Norway spruce cutting clones on different sites. Silva Fenn 40:247256 Raymond CA, Dickson R, Rowell D, Blakemore P, Clark N, Williams M, Freischmidt G, Joe B (2004) Wood and ber properties of dry-land conifers. RIRDC Publication No 04/099, p 69 SAS Institute (1990) SAS/STAT Users Guide. Version 6, 4th edn, SAS, Cary Savva Y, Koubaa A, Tremblay F, Bergeron Y (2010) Effects of radial growth, tree age, climate, and seed origin on wood density of diverse jack pine populations. Trees 24:5365 Smith DM (1954) Maximum moisture content method for determining specic gravity of small wood samples. US Forest Service, Forest products Lab. Report No:2014 Tassisa G, Burkhart HE (1997) Juvenilemature wood demarcation in loblolly pine trees. Wood Fiber Sci 30:119127 Usta HZ (1991) A study on the yield of Pinus brutia Ten. Plantations. Turkish Forest Research Institute, Technical Bulletin No: 219, Ankara, Turkey, p 138 Weinstein A (1989) Geographic variation and phenology of Pinus halepensis, P. brutia and P. eldarica in Israel. For Ecol Manag 27:99108 Zobel BJ, Sprague JR (1998) Juvenile wood in forest trees. Springer Series in Wood Science, Springer-Verlag Zobel BJ, Van Buijtenen JP (1989) Wood Variationits causes and control. Springer Series in Wood Science, Springer-Verlag, p 363

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Trees
Structure and Function
Editors-in-Chief: U. Lttge; R.D. Guy

Covers physiology, biochemistry, functional anatomy, structure and ecology of trees and other woody plants Also presents research articles on pathology and technological problems that contribute to basic understanding of tree structure and function Publishes reviews on selected topics
Trees - Structure and Function publishes original papers on the physiology, biochemistry, functional anatomy, structure and ecology of trees and other woody plants. Also presented are articles concerned with pathology and technological problems, when they contribute to the basic understanding of structure and function of trees. In addition to original papers and short communications, the journal publishes reviews on selected topics concerning the structure and function of trees. The Founding Editor of Trees - Structure and Function is Hubert Ziegler. The Editors-in-Chief are Robert D. Guy, Department of Forest Sciences, University of British Columbia, Vancouver, Canada, and Ulrich E. Lttge, Botanisches Institut der TU, Darmstadt, Germany. 5-Year Impact Factor:1.900 (2010)* Impact Factor: 1.444 (2010), Journal Citation Reports, Thomson Reuters On the homepage of Trees at springer.com you can
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Editorial Board Founding Editor Hubert Ziegler Managing Editors for papers from the Americas: Robert D. Guy The University of British Columbia Department of Forest Sciences Faculty of Forestry # 3041-2424 Main Mall Vancouver BC V6T 1Z4, Canada e-mail: guy@interchange.ubc.ca for papers from all other countries: Ulrich E. Lttge Botanisches Institut der TU Schnittspahnstrasse 3-5 64287 Darmstadt, Germany e-mail: luettge@bio.tu-darmstadt.de Tel.: +49-6151-163200 Fax: +49-6151-164630 Editorial Assistant Verena Kastrup Botanisches Institut der TU Schnittspahnstrasse 3-5 64287 Darmstadt, Germany Editors Mark Adams Faculty of Agriculture Food and Natural Resources University of Sydney Sydney, NSW 2006 Australia e-mail: mark.adams@usyd.edu.au Roni Aloni Tel Aviv University Department of Plant Sciences 69978 Tel Aviv, Israel e-mail: alonir@post.tau.ac.il Erwin Beck Lehrstuhl Pflanzenphysiologie Universitt Bayreuth Universittsstrasse 30 95440 Bayreuth, Germany e-mail: erwin.beck@uni-bayreuth.de Wolfgang Bilger Botanisches Institut Abt. kophysiologie der Pflanzen Am Botanischen Garten 3-9

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24118 Kiel, Germany e-mail: wbilger@bot.uni-kiel.de Marcos Buckeridge Department of Botany Institute of Biosciences Rua do Matao, 277 Sao Paulo, SP Brazil PO Box 11461 Postal Code 05422-970 e-mail: msbuck@usp.br Thomas Buckley School of Biological, Earth and Environmental Science UNSW Sydney NSW 2052, Australia e-mail: Thomas.Buckley@ensisjv.com Francisco M. Cnovas Unidad asociada UMA-CSIC Universidad de Mlaga Campus de Teatinos 29071 Mlaga, Spain e-mail: canovas@uma.es John E. Carlson The School of Forest Resources Pennsylvania State University 323 Forest Resources Building University Park, PA 16802, USA e-mail: jec16@psu.edu Herv?Cochard UMR - Physiologie Intgrede l'Arbre Frutier et Forestier INRA, Site de Crouelle 63039 Clermont-Ferrand, France e-mail: cochard@clermont.inra.fr Bettina Engelbrecht Department of Biology San Francisco State University 1600 Holoway Ave San Francisco, CA 94132, USA e-mail: bettina.engelbrecht@gmail.com Thierry Fourcaud UMR AMAP botAnique et bioinforMatique de l'Architecture des Plantes TA A-51/PS2 (Bat. PSII, Bur. 105) Boulevard de la Lironde 34398 Montpellier Cedex 5, France e-mail: thierry.fourcaud@cirad.fr Arthur Geler University of Freiburg Core Facility Metabolomics Centre for System Biology (ZBSA) Habsburgerstr. 49

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79104 Freiburg, Germany e-mail: arthur.gessler@sonne.uni-freiburg.de Thorsten Grams Technische Universitt Mnchen Department of Ecology, Ecophysiology of Plants Am Hochanger 13 85354 Freising, Germany e-mail: grams@wzw.tum.de Rdiger Hampp Universitt Tbingen Botanisches Institut Auf der Morgenstelle 1 72076 Tbingen, Germany e-mail: ruediger.hampp@uni-tuebingen.de Taizo Hogetsu The University of Tokyo Asian Natural Science Center Midori-cho 1-1-8 Nishtokyo-shi Tokyo 188-0002, Japan e-mail: ho@fr.a.u-tokyo.ac.jp Hamlyn G. Jones Plant Research Unit Div. Environmental and Applied Biology University of Dundee at SCRI Dundee DD2 5DA, U.K. e-mail: h.g.jones@dundee.ac.uk Olavi Junttila University of Troms?BR>9037 Troms? Norway olavi.junttila@ib.uit.no Steven W. Leavitt University of Arizona Laboratory of Tree-Ring Research Tucson, AZ 85721, USA e-mail: sleavitt@ltrr.arizona.edu Sune Linder Southern Swedish Forest Research Centre Swedish University of Agricultural Sciences P.O. Box 49 230 53 Alnarp, Sweden e-mail: sune.linder@ess.slu.se Catherine Lovelock Centre for Marine Studies University of Queensland St Lucia QLD 4072, Australia e-mail: c.lovelock@uq.edu.au John E. Major Canadian Forest Service Atlantic Forestry Centre PO Box 4000, Room 3-506

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Fredericton NB, E3B 5P7, Canada jmajor@nrcan.gc.ca Rainer Matyssek Lehrstuhl fr Forstbotanik der Universitt Mnchen Am Hochanger 3 85354 Freising, Germany e-mail: matyssek@bot.forst.tu-muenchen.de Wolfgang Owald WZW - Department fr kologie Am Hochanger 3 85375 Freising, Germany e-mail: osswald@wzw.tum.de Hardy Pfanz Institut fr Angewandte Botanik Universitt Duisburg-Essen Campus Essen Universittsstr. 5 45117 Essen, Germany e-mail: hardy.pfanz@uni-essen.de Heinz Rennenberg Institut fr Forstbotanik und Baumphysiologie Universitt Freiburg Am Flughafen 17 79085 Freiburg, Germany e-mail: here@uni-freiburg.de Thomas Speck Universitt Freiburg Institut fr Biologie II/III Schnzlestr. 1 79104 Freiburg, Germany e-mail: thomas.speck@biologie.uni-freiburg.de Keiji Takabe Kyoto University Laboratory of Structure of Plant Cells Division of Forest and Biomaterial Sciences Graduate School of Agriculture Kyoto 606-8502, Japan e-mail: kjtakabe@kais.kyoto-u.ac.jp Dieter Treutter Technische Universitt Mnchen Fachgebiet Obstbau Alte Akademie 16 85350 Freising, Germany e-mail: dieter.treutter@wzw.tum.de Klaus Winter Smithsonian Tropical Institute P.O. Box 2072 Balboa, Rep. Panama e-mail: winterk@tivoli.si.edu

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Maciej A. Zwieniecki The Arnold Arboretum of Harvard University Biological Laboratories 16 Divinity Ave. Cambridge, MA 02138, USA e-mail: mzwienie@oeb.harvard.edu

Hubert Ziegler Managing Editors for papers from the Americas: Robert D. Guy The University of British Columbia Department of Forest Sciences Faculty of Forestry # 3041-2424 Main Mall Vancouver BC V6T 1Z4, Canada e-mail: guy@interchange.ubc.ca for papers from all other countries: Ulrich E. Lttge Botanisches Institut der TU Schnittspahnstrasse 3-5 64287 Darmstadt, Germany e-mail: luettge@bio.tu-darmstadt.de Tel.: +49-6151-163200 Fax: +49-6151-164630 Editorial Assistant Verena Kastrup Botanisches Institut der TU Schnittspahnstrasse 3-5 64287 Darmstadt, Germany Editors Mark Adams Faculty of Agriculture Food and Natural Resources University of Sydney Sydney, NSW 2006 Australia e-mail: mark.adams@usyd.edu.au Roni Aloni Tel Aviv University Department of Plant Sciences 69978 Tel Aviv, Israel e-mail: alonir@post.tau.ac.il Erwin Beck Lehrstuhl Pflanzenphysiologie

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Universitt Bayreuth Universittsstrasse 30 95440 Bayreuth, Germany e-mail: erwin.beck@uni-bayreuth.de Wolfgang Bilger Botanisches Institut Abt. kophysiologie der Pflanzen Am Botanischen Garten 3-9 24118 Kiel, Germany e-mail: wbilger@bot.uni-kiel.de Marcos Buckeridge Department of Botany Institute of Biosciences Rua do Matao, 277 Sao Paulo, SP Brazil PO Box 11461 Postal Code 05422-970 e-mail: msbuck@usp.br Thomas Buckley School of Biological, Earth and Environmental Science UNSW Sydney NSW 2052, Australia e-mail: Thomas.Buckley@ensisjv.com Francisco M. Cnovas Unidad asociada UMA-CSIC Universidad de Mlaga Campus de Teatinos 29071 Mlaga, Spain e-mail: canovas@uma.es John E. Carlson The School of Forest Resources Pennsylvania State University 323 Forest Resources Building University Park, PA 16802, USA e-mail: jec16@psu.edu Herv?Cochard UMR - Physiologie Intgrede l'Arbre Frutier et Forestier INRA, Site de Crouelle 63039 Clermont-Ferrand, France e-mail: cochard@clermont.inra.fr Bettina Engelbrecht Department of Biology San Francisco State University 1600 Holoway Ave San Francisco, CA 94132, USA e-mail: bettina.engelbrecht@gmail.com

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Thierry Fourcaud UMR AMAP botAnique et bioinforMatique de l'Architecture des Plantes TA A-51/PS2 (Bat. PSII, Bur. 105) Boulevard de la Lironde 34398 Montpellier Cedex 5, France e-mail: thierry.fourcaud@cirad.fr Arthur Geler University of Freiburg Core Facility Metabolomics Centre for System Biology (ZBSA) Habsburgerstr. 49 79104 Freiburg, Germany e-mail: arthur.gessler@sonne.uni-freiburg.de Thorsten Grams Technische Universitt Mnchen Department of Ecology, Ecophysiology of Plants Am Hochanger 13 85354 Freising, Germany e-mail: grams@wzw.tum.de Rdiger Hampp Universitt Tbingen Botanisches Institut Auf der Morgenstelle 1 72076 Tbingen, Germany e-mail: ruediger.hampp@uni-tuebingen.de Taizo Hogetsu The University of Tokyo Asian Natural Science Center Midori-cho 1-1-8 Nishtokyo-shi Tokyo 188-0002, Japan e-mail: ho@fr.a.u-tokyo.ac.jp Hamlyn G. Jones Plant Research Unit Div. Environmental and Applied Biology University of Dundee at SCRI Dundee DD2 5DA, U.K. e-mail: h.g.jones@dundee.ac.uk Olavi Junttila University of Troms?BR>9037 Troms? Norway olavi.junttila@ib.uit.no Steven W. Leavitt University of Arizona Laboratory of Tree-Ring Research Tucson, AZ 85721, USA e-mail: sleavitt@ltrr.arizona.edu

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Sune Linder Southern Swedish Forest Research Centre Swedish University of Agricultural Sciences P.O. Box 49 230 53 Alnarp, Sweden e-mail: sune.linder@ess.slu.se Catherine Lovelock Centre for Marine Studies University of Queensland St Lucia QLD 4072, Australia e-mail: c.lovelock@uq.edu.au John E. Major Canadian Forest Service Atlantic Forestry Centre PO Box 4000, Room 3-506 Fredericton NB, E3B 5P7, Canada jmajor@nrcan.gc.ca Rainer Matyssek Lehrstuhl fr Forstbotanik der Universitt Mnchen Am Hochanger 3 85354 Freising, Germany e-mail: matyssek@bot.forst.tu-muenchen.de Wolfgang Owald WZW - Department fr kologie Am Hochanger 3 85375 Freising, Germany e-mail: osswald@wzw.tum.de Hardy Pfanz Institut fr Angewandte Botanik Universitt Duisburg-Essen Campus Essen Universittsstr. 5 45117 Essen, Germany e-mail: hardy.pfanz@uni-essen.de Heinz Rennenberg Institut fr Forstbotanik und Baumphysiologie Universitt Freiburg Am Flughafen 17 79085 Freiburg, Germany e-mail: here@uni-freiburg.de Thomas Speck Universitt Freiburg Institut fr Biologie II/III Schnzlestr. 1 79104 Freiburg, Germany e-mail: thomas.speck@biologie.uni-freiburg.de

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Keiji Takabe Kyoto University Laboratory of Structure of Plant Cells Division of Forest and Biomaterial Sciences Graduate School of Agriculture Kyoto 606-8502, Japan e-mail: kjtakabe@kais.kyoto-u.ac.jp Dieter Treutter Technische Universitt Mnchen Fachgebiet Obstbau Alte Akademie 16 85350 Freising, Germany e-mail: dieter.treutter@wzw.tum.de Klaus Winter Smithsonian Tropical Institute P.O. Box 2072 Balboa, Rep. Panama e-mail: winterk@tivoli.si.edu Maciej A. Zwieniecki The Arnold Arboretum of Harvard University Biological Laboratories 16 Divinity Ave. Cambridge, MA 02138, USA e-mail: mzwienie@oeb.harvard.edu

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Thomson Reuters Master Journal List JOURNAL LIST Search terms: TREES Total journals found: 1 1. TREES-STRUCTURE AND FUNCTION Bimonthly ISSN: 0931-1890 SPRINGER, 233 SPRING ST, NEW YORK, USA, NY, 10013 1. Science Citation Index 2. Science Citation Index Expanded 3. Current Contents - Agriculture, Biology & Environmental Sciences 4. BIOSIS Previews

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12.04.2012