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Scientific Area A Plant structure, function and development

59

Variations in quantitative vessel element characters of Cerasus avium (Rosaceae) in Turkey


Barbaros Yaman
Department of Forest Botany, Faculty of Forestry, Zonguldak Karaelmas University, 74100 Bartin, Turkey, e-mail: yamanbar2000@yahoo.com

Abstract.

Variations in quantitative vessel element characters of the wood of Cerasus avium were investigated in relation to altitude and tree-ring width. According to multiple regression analysis using altitude and tree-ring width as independent variables, significant correlations were obtained at the 0.001 level for vessel density and vulnerability ratio, at the 0.01 level for mesomorphy ratio, and at the 0.05 level for vessel tangential diameter and element length. The number of vessels per group does not show any correlation with independent variables. Non-anatomical factors (altitude and tree-ring width) explain 25.450.1 % of variation for dependent variables. altitude, Cerasus avium, ecology, tree-rings, wood anatomy

Key words:

Introduction
Cerasus avium (L.) Moench is one of the native taxa in Turkey; it is distributed from sea level up to 1600 m alt. in deciduous forests of Euxine province (Browicz 1972). Euxine province, one of the two sub-provinces of EuroSiberian Region, lies between Melet River in Ordu and Istranca Mt in Krklareli along Black Sea Region of Turkey (Yaltrk & Efe 1989) (Fig. 1). In general, there is a humid climate type in this subprovince (Erin 1996). The basic and detailed studies in relation to ecological wood anatomy were carried out by different researchers (Carlquist 1966, 1975, 1977a, b, 1980; Baas 1973, 1976; Van der Graaff & Baas 1974; Baas & al. 1983; Carlquist & Hoekman 1985; Fahn & al.

Fig. 1. Euxine province between arrows, and localities of wild cherry trees sampled in this province of Turkey ().

1986). In recent years, many studies have been realized on this subject in different regions of the world (Zhang & al. 1992; Lindorf 1994, 1997; Noshiro & al. 1994, 1995; Noshiro & Suzuki 1995; Villar-Salvador & al. 1997; Mauseth 1999; Alves & Angyalossy-Alfonso 2000, 2002; Cooper & Cass 2001). Most of the studies mentioned above indicated some ecological trends in vessel element characters. It has been explained by Carlquist (1975) and examined by Baas (1976) in detail that anatomical diversity in wood is a function of selective pressures of environmental conditions such as water availability, temperature, etc. Two ratios based on vessel diameter, vessel density and vessel element length were formulated by Carlquist (1977b) and Carlquist & DeBuhr (1977) for ecological interpretation in wood anatomy: vulnerability (mean vessel diameter divided by mean vessel density) and mesomorphy (vulnerability multiplied by vessel element length). In his book, entitled "Comparative Wood Anatomy", Carlquist (1988) evaluated the significance of vulnerability and mesomorphy equations as compared to Hagen-Poiseuille equation in detail. As for Turkey, there are a small number of studies related to ecological wood anatomy. At first, Yaltrk (1968) studied anatomical characteristics of Turkish maples wood with relation to the humidity of

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Plant, fungal and habitat diversity investigation and conservation Proceedings of IV BBC Sofia ' 2006

the sites. anl (1977) examined anatomical features of beech woods related to altitude in the different regions of Turkey. After then, some studies mentioning the relationships between wood anatomical characters and ecology have been carried out in Turkey (Gerek & al. 1998; Merev & Yavuz 2000; Yaman 2002; Serdar 2003). Because there have been a great variety of woody taxa and their habitats in Turkey, and a small number of these taxa have been studied so far in relation to ecological wood anatomy, Turkey has an important potential for this research area. In the present study, the wood of C. avium was selected as research material because its altitudinal range in Euxine province is enough to investigate altitudinal trends. The variations in quantitative vessel element characters, vulnerability and mesomorphy ratios of the wood of this species were examined in relation to altitude and tree-ring width.

mined from macerated materials and measured from tip to tip (including the element tails). For quantification of vessel grouping, the method proposed by Carlquist (1988) was used. Twenty-five measurements were performed for the mean of each quantitative anatomical character. In addition, increment core was extracted by using increment borer from breast height of trunk for each sample tree. Tree-ring widths belonging to last ten years were measured on the increment cores, and later their mean was calculated for each tree. Data gathered on altitude, tree-ring width, selected anatomical characters, vulnerability and mesomorphy ratios were analysed by correlation and multiple regression methods in the SPSS 9.0 program. In the multiple regression analysis, altitude and tree-ring width were considered as independent variables, and the selected wood anatomical characters, vulnerability and mesomorphy ratios were used as dependent variables.

Material and methods


The mature wood specimens were extracted from 24 wild cherry trees located at different altitudes of Euxine subprovince (Table 1). During sampling, the trees with even-aged (about 50 years old) were preferred for minimizing the effect of tree stem diameter and age on quantitative anatomical characters of wood. The wood specimens were extracted from the same side (north) at breast height of trunks and later they were given shape to cubic. Prior to sectioning, the cubicshaped woods were boiled, and later they were put in the solution of "glycerine waterethyl alcohol" during several days for softening. After these procedures, the transverse, tangential and radial sections were taken from cubic-shaped woods by using a Euromex sliding microtome. For preparation of wood sections and macerations, standard procedures were applied (Yaltrk 1971). Vessel diameter, vessel density (the number of vessels per mm2) and the number of vessels per group were determined in transverse sections. Vessel diameter was measured based on lumen. Vessel element length was deterTable 1. Some information for the localities of the selected sample trees of C. avium in Euxine province of Turkey. RFD Giresun Giresun Amasya Sinop Zonguldak Zonguldak Zonguldak Zonguldak Zonguldak Zonguldak Zonguldak Zonguldak Zonguldak Zonguldak Bolu Bolu Bolu Bolu Bolu Bolu Adapazari Istanbul Istanbul Istanbul FE Ordu Unye Samsun Ayancik Yenice Yenice Yenice Yenice Yenice Ulus Ulus Bartin Bartin Dirgine Akcakoca Akcakoca Akcakoca Duzce Duzce Duzce Izmit Demirkoy Demirkoy Demirkoy FTD Merkez Merkez Ayvacik Goldagi Kavakli Kavakli Kavakli Kavakli Kavakli Drahna Drahna Yenihan Yenihan Kozdere Altincay Altincay Altincay Cicekli Cicekli Merkez Dilovasi Macara Kadinkule Kadinkule CST GOM GUM ASA SAG ZYK1A ZYK1B ZYK2 ZYK3A ZYK3B ZUD1 ZUD2 ZBY1 ZBY2 ZDK BAA1 BAA2 BAA3 BDC1 BDC2 BDM AID IDM IDK1 IDK2 LFCN Kursuncali Cataltepe Senpinar Cinarduzu 4 4 21 28 28 Nun deresi Nun deresi Kirazlik Kirazlik Elemen Altincay Altincay Altincay Yanik Yanik Merkez Radar Pirgoplu Kadinkule Velika ALT 980 180 210 580 410 410 790 1350 1350 890 1070 480 480 1150 280 320 340 940 940 220 480 180 650 470 EXP North North Northwest Northwest North North North Northeast Northeast Northeast Northeast Northeast Northeast North North Northeast Northeast Northwest Northwest North North Northeast North North DT 19.06.2001 20.06.2001 20.06.2001 21.06.2001 10.08.2000 10.08.2000 10.08.2000 10.08.2000 10.08.2000 18.09.2000 18.09.2000 04.10.2000 04.10.2000 01.09.2001 06.10.2000 06.10.2000 06.10.2000 04.10.2001 04.10.2001 04.10.2001 03.10.2001 28.06.2001 29.06.2001 29.06.2001

Abbreviations: RFD Regional forest directorate; FE Forest enterprise; FTD Forest territorial division; CST Code of sample tree; LFCN Locality or forest compartment number; ALT Altitude; EXP Exposure; DT Date of obtaining of wood specimen.

Scientific Area A Plant structure, function and development

61

Results
Cerasus avium has a wood with semi-ring porous; however, it tends to diffuse porous in narrower rings. Tree-rings are distinct because of size differences in vessel diameter between latewood and earlywood in successive tree-rings, and flattened fibres at the end of latewood. Mean vessel tangential diameter is 53.61 m (range 47.1963.15 m). Vessel density shows a range from 80.70 to 159.65; and its mean is 125.01. Mean vessel element length is 399.65 m (range 346448.40 m). The number of vessels per group ranges from 1.34 to 2.01 on average, and its mean is 1.67. Vessels are solitary, or in radial multiples or clusters. Solitary vessels are oval to round, and grouped ones are almost angular in outline. There are well-developed helical thickenings on vessel wall, and vessel elements have simple perforation plates. The means of vulnerability and mesomorphy ratios for C. avium are respectively 0.4458 and 178.08 in Euxine province. Vulnerability and mesomorphy ratios, and the quantitative data belonging to selected anatomical characters are given separately for each sample tree in Table 2. According to Pearson correlation analysis (Table 3), the correlation between two non-anatomical features (altitude and tree-ring width) is not statistically significant. However, vessel tangential diameter and vessel density show a significant correlation with altitude (correlation coefficients 0.514 and 0.463, respectively). There is no significant correlation between altitude and other two selected anatomical characters. In other words, while decreasing vessel diameter of C. avium wood from low to high altitudes in Euxine province, vessel density increases; however, there is not any significant relationship between vessel element length, the number of vessels per group and altitude. While only vessel density has a significant correlation with tree-ring width, other three anatomical characters do not show a relationship with this non-anatomical feature. The ratios of vulnerability and mesomorphy based on vessel diameter, vessel density and element length have significant correlations with both altitude and tree-ring width. In other words, vulnerability and mesomorphy ratios estimated for C. avium decrease from low to high altitude, and increase from narrow tree-rings to wider ones.

Table 2. Non-anatomical and selected anatomical characters of C. avium wood. CST GUM IDM ASA BDM BAA1 BAA2 BAA3 ZYK1A ZYK1B IDK2 ZBY1 ZBY2 AID SAG IDK1 ZYK2 ZUD1 BDC1 BDC2 GOM ZUD2 ZDK ZYK3B ZYK3A MEAN S.D. ALT TRW VTD VMM2 VEL VGR VUL (m) (mm) (m) (m) 180 180 210 220 280 320 340 410 410 470 480 480 480 580 650 790 890 940 940 980 1070 1150 1350 1350 5.24 1.95 4.47 6.11 5.47 8.78 3.01 2.75 2.49 4.28 2.89 3.01 1.30 7.41 0.59 3.36 1.20 7.50 1.95 1.59 2.08 3.64 3.54 2.88 3.64 2.14 59.69 54.37 53.59 51.49 52.32 63.15 55.66 55.43 54.50 54.93 53.99 53.76 51.92 48.38 54.11 55.86 54.70 54.27 52.04 54.63 48.00 52.10 50.60 47.19 53.61 3.42 80.70 151.70 106.40 83.15 126.70 97.20 128.75 129.30 134.20 134.80 112.40 119.20 142.35 101.00 154.60 148.20 109.25 107.60 118.20 120.80 145.60 134.00 154.60 159.65 125.01 22.30 395.20 416.40 408.40 346.00 446.40 424.40 405.20 431.20 445.20 394.00 374.40 448.40 403.60 346.80 403.20 407.20 430.80 363.60 420.80 394.40 361.60 364.00 404.80 355.60 399.65 31.28 1.34 1.67 1.63 1.62 1.75 1.70 1.87 1.85 1.63 1.70 1.58 1.58 1.64 1.41 1.73 1.96 1.70 1.72 1.52 1.70 1.63 1.56 1.70 2.01 1.67 0.15 0.7396 0.3584 0.5036 0.6192 0.4129 0.6497 0.4323 0.4287 0.4061 0.4075 0.4803 0.4510 0.3647 0.4790 0.3500 0.3769 0.5007 0.5044 0.4403 0.4522 0.3296 0.3888 0.3273 0.2956 0.4458 0.1055 MESO 292.29 149.23 205.68 214.24 184.34 275.73 175.17 184.85 180.78 160.54 179.84 202.23 147.19 166.10 141.12 153.48 215.70 183.39 185.27 178.35 119.20 141.51 132.49 105.10 178.08 43.36

Abbreviations: CST Code of sample tree; ALT Altitude; TRW Tree-ring width; VTD Vessel tangential diameter; VMM2 Vessel density (the number of vessels per mm2); VEL Vessel element length; VGR The number of vessels per group; VUL Vulnerability ratio; MESO Mesomorphy ratio; S.D. Standard deviation. Table 3. Correlations of non-anatomical and selected wood anatomical characters, vulnerability and mesomorphy ratios of C. avium. ALT ALT TRW VTD VMM2 VEL VGR VUL MESO 1.000 -0.260 -0.514* 0.463* -0.341 0.253 -0.520** -0.584** TRW VTD VMM2 VEL VGR VUL MESO

1.000 0.254 -0.609** -0.270 -0.207 0.599** 0.484*

1.000 -0.396 0.475* -0.040 0.619** 0.754***

1.000 0.114 0.575** -0.939*** -0.851***

1.000 0.124 1.000 -0.025 -0.501* 1.000 0.302 -0.427* 0.943** 1.000

*** = significant at the 0.001 level; ** = significant at the 0.01 level; * = significant at the 0.05 level. Abbreviations: as in Table 2.

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Plant, fungal and habitat diversity investigation and conservation Proceedings of IV BBC Sofia ' 2006

In addition, the selected quantitative anatomical features, vulnerability and mesomorphy ratios have also correlations with one another. Vessel tangential diameter is positively correlated with vessel element length, vulnerability and mesomorphy ratios. Vessel density is positively correlated with the number of vessels per group, and negatively correlated with vulnerability and mesomorphy ratios. Besides, the number of vessels per group shows a negative correlation with vulnerability and mesomorphy ratios. According to the multiple regression analysis using altitude and tree-ring width as independent variables, significant correlations were obtained at the 0.001 level for vessel density and vulnerability ratio, at the 0.01 level for mesomorphy ratio, and at the 0.05 level for vessel tangential diameter and element length (Table 4). The number of vessels per group does not show any correlation with independent variables. Determination coefficients (R2) are 0.2540.501 for significant dependent vari180 160 Vessel Density 140 120 100 80 60 40 20 0 0 2 4 6 8 10 Tree-Ring Width

Table 4. Multiple regression analysis of selected anatomical characters, vulnerability, and mesomorphy ratios. DV R R2 F-ratio Partial Regression Coefficient Constant VTD
2

Standardized Regression Coefficient ALT TRW

ALT -0.0044
*

TRW

0.529 0.280 4.077

55.667 132.452 1.659 0.427

0.2060 -0.481 0.129

VMM 0.686 0.470 9.309 VEL VGR VUL

***

0.0196 -5.4460** 0.327 -0.524 0.0000 -0.0107 0.214 -0.152 -0.0001* 0.0244** -0.390 0.498

0.504 0.254 3.570* 0.293 0.086 0.983 0.708 0.501 10.536


***

443.567 -0.0372* -5.6030 -0.440 -0.384

MESO 0.678 0.460 8.930**

188.077 -0.0575** 7.2140* -0.491 0.357

*** = significant at the 0.001 level; ** = significant at the 0.01 level; * = significant at the 0.05 level. Abbreviations: DV Dependent variables; R Multiple correlation coefficient; R2 Coefficient of determination; other abbreviations as in Table 2.

ables. In other words, non-anatomical factors (ALT and TRW) explain 25.450.1 % of total variation for significant dependent variables. For altitude, standardized regression coefficients are significant at the 0.01 level for mesomorphy ratio, and at the 0.05 level for vessel tangential diameter, vessel element length and vulnerability ratio. For tree-ring width, they are significant at the 0.01 level for vessel density and vulnerability ratio, and at the 0.05 level for mesomorphy ratio. The greatest standardized regression coefficients between dependent variables and non-anatomical factors are 0.524 for vessel density, 0.498 for vulnerability and 0.491 for mesomorphy. Change in the values of these dependent variables in respect to independent variables is plotted in Figs 24.
350 Mesomorphy Ratio 300 250 200 150 100 50 0 0 500 Altitude 1000 1500

Fig. 2. Trend between vessel density and tree-ring width.


0.8 0.7 Vulnerability Ratio 0.6 0.5 0.4 0.3 0.2 0.1 0 0 2 4 6 8 10 Tree-Ring Width

Fig. 3. Trend between vulnerability ratio and tree-ring width.

Fig. 4. Trend between mesomorphy ratio and altitude.

Scientific Area A Plant structure, function and development

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Discussion
Wood anatomy of Rosaceae family in China was studied by Zhang and Baas (1992), and wood structure of this family in relation to ecology, habit and phenology was evaluated in details by Zhang & al. (1992). In these articles, there were also one variety and four different samples of C. avium among the Rosaceae species studied. The mean values of some quantitative vessel characters in the woods of four different specimens of C. avium in China are 4855 m for vessel tangential diameter, 90176 vessels per mm2 for vessel density and 300390 m for vessel element length. The mean values in wood of C. avium var. decumana for the aforementioned vessel characters were 50 m, 220 vessels per mm2, and 390 m, respectively (Zhang & Baas 1992). As for Turkish C. avium examined in the present study, the mean values are 53.61 m for vessel tangential diameter, 125.01 for vessel density and 399.65 m for vessel element length. According to IAWA Committee (1989), it may be classified as number 41 for vessel tangential diameter, number 50 for vessel density and number 53 for vessel element length. Altitude has a significant effect on vessel tangential diameter, vessel element length, mesomorphy and vulnerability ratios of C. avium wood. While vessel tangential diameter and vessel element length tend to decrease with increasing of altitude in Euxine province, vulnerability and mesomorphy ratios tend to be lower values. Carlquist (1977b) introduced the concept of vulnerability to wood anatomists with regard to embolism of secondary xylem. Very narrow vessel diameter corresponds with low vulnerability values (<1), which reflect high conductive safety, while high vulnerability values (>3) are indicative of taxa having high conductive efficiency. The mean vulnerability value of C. avium in Euxine province is 0.4458. According to this vulnerability value, it can be explained that the wood of C. avium in this province shows high conductive safety. Furthermore, helical thickenings on vessel wall of C. avium may be considered as a feature raising conductive safety. Carlquist (1988) pointed out that the correlation between some quantitative vessel element features and altitude should be connected with factors of water availability and temperature, since altitude is not an ecological factor in itself. In Euxine province, while annual precipitation increases from sea level up to high altitude, mean annual temperature decreases. Among localities of sample trees, the amount of annual precip-

itation ranges from 764.30 mm up to 1583 mm, and mean annual temperature has a range between 7.15 C and 13.48 C (Yaman 2002). In spite of enough precipitation in Euxine province (humid climate), low temperatures may limit growth of trees at high elevation. Low temperatures at high elevations affect the viscosity of water both in soil and in stem (epel 1993), thus taking water up from soil and transport of water in secondary xylem may be negatively affected. The change in quantitative characters of vessel elements in C. avium associated with changing temperature along altitudinal gradients was consistent with the hypothesis of Roderick & Berry (2001). These authors pointed out that the effect of temperature on the viscosity of water will affect vessel size and / or number and therefore wood density. Recently, the physiological link between water viscosity and wood anatomy was examined in Eucalyptus camaldulensis by Thomas & al. (2004). Carlquist (1988) indicated that narrower vessel diameter and shorter vessel element length are related to conductive safety. Becoming narrower of vessel diameter and shorter of vessel element length of C. avium wood growing in high altitudes can be explained as an indication of conductive safety. Decreasing of mesomorphy value from low up to high altitudes shows that high elevations in Euxine province (despite not xeric sites) cause a xeromorphic effect on the wood of C. avium because of the low temperature. Mean mesomorphy value of C. avium in this province is 178.08. In general, this species in Euxine province is mesomorphic with respect to its wood anatomy; however, it tends to have xeromorphic wood features (narrower vessel diameter and shorter vessel element length) at high altitudes of this province as if it is in xeric site. As known, it was indicated that low values of mesomorphy ratio (<100) show xeromorphy, and its high values show mesomorphy with respect to wood anatomy (Carlquist 1977b). There is not a significant correlation between altitude and tree-ring width in this research; however, tree-rings width significantly affects vessel density, vulnerability and mesomorphy ratios. Vessel density increases with narrowing of tree-rings in the wood of C. avium in Euxine province. This may be explained as the high altitudinal conditions that cause the narrowing of tree-rings also cause increase in vessel density. Positive correlations between vulnerability, mesomorphy and tree-ring width seem to be quite logical results when environmental conditions causing the narrowing of tree-ring are considered.

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