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SEEFOR 6 (1): 15-23
Article ID: 46
DOI: http://dx.doi.org/10.15177/seefor.15-03

Original scientific paper


The Effects of Exposure, Elevation and Tree Age on Seed Characteristics of Fagus orientalis Lipsky

Murat Ertekin 1, Erol Kırdar 2, Sezgin Ayan 3*

1 Necmettin Erbakan University, Department of Crop and Animal Production, TR-42370 Seydişehir, Konya, Turkey
2 Bartin University, Faculty of Forestry, Department of Silviculture, TR-74100 Bartın, Turkey
3 Kastamonu University, Faculty of Forestry, Department of Silviculture, TR-37100 Kuzeykent, Kastamonu, Turkey

* Corresponding author: e-mail: 

ERTEKIN M, KIRDAR E, AYAN S 2015 The Effects of Exposure, Elevation and Tree Age on Seed Characteristics of Fagus orientalis Lipsky. South-east Eur for 6 (1): 15-23. DOI: http://dx.doi.org/10.15177/seefor.15-03

Received: 18 Jan 2015 / Accepted: 24 Feb 2015 / Published online: 16 Mar 2015

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Background and Purpose: Natural or artificial regeneration, rehabilitation, and conversion from coppice to high forest are important practices in Oriental beech (Fagus orientalis Lipsky) forests in Turkey. Studies of the seeds of this species have increased in number because mast years are infrequent and seed germination is inhibited by dormancy. In this study we quantified the effects of tree age (40-59, 60-79 and 80-99 years), stand exposure (north, west, east and south) and elevation (600 and 800 m a.s.l.) on seed characteristics (germination, moisture content, and weight) of Oriental beech.
Material and Methods: The seeds used in this study were collected from natural beech forest at Kumluca, Bartin, in the western Black Sea region of Turkey. Experiments were carried out in the laboratory and the greenhouse of Bartin University.
Results: Seed germination and moisture content varied significantly by elevation, and seed germination was strongly influenced by elevation. Moisture content was 14% at 600 m and 16% at 800 m. The effects of elevation and tree age on 100 seed weights were not significant but exposure had a significant effect. The highest 100 seed weight was recorded for trees on southern exposures and the highest germination percentage of 82% was recorded for trees on northern exposures.
Conclusions: In conclusion, since oriental beech seedlings are produced by generative propagation method, seeds should be harvested in optimum distribution area of beech, from average ages and phenotypically plus tree.

Keywords: Oriental beech, natural forest, tree age, elevation, exposure, seed characteristics 



One of the most abundant and economically important hardwood genera in northern hemisphere temperate forests is Fagus [1]. In Turkey, Oriental beech (Fagus orientalis Lipsky) is a common hardwood tree species that regenerates naturally in Turkish forests where species diversity is rich due to the variety of growing conditions. It is a shade-tolerant species, and optimum growth conditions for Oriental beech are on the north-facing slopes of the North Anatolian orogenic belt in the Black Sea Region and the Istranca Mountains in Thrace. Oriental beech forests in Turkey cover 1810079 ha, composing nearly 8.5% of the country’s total forest area [2]. The elevation range of Oriental beech in the Balkans is 10-800 m above sea level (m a.s.l.). In Turkey, it grows at 1500-1700 m a.s.l. in Black Sea inner-valleys and up to 2000 m a.s.l. in the Aegean mountains [3], [4], [5], [6], [7]. Most Oriental beech forests are distributed in northern Turkey. However, relict Oriental beech forests are distributed in the Eastern Mediterranean region of Turkey, including the provinces of Adana, Osmaniye, Hatay, and Kahramanmaras [8]. Oriental beech forests in Turkey usually have mixed structure with Nordmann fir (Abies nordmanniana (Steven) Spach, Uludag fir (A. nordmanniana (Stev.) Spach. subsp. bornmuelleriana (Mattf.) Coode&Cullen), Kazdagı fir (A. equi-trojani Aschers et Sinten), Scotch pine (Pinus sylvestrisL.), Anatolian black pine (P. nigra Arnold subsp. pallasiana (Lamb.) Holmboe) and Oriental spruce (Picea orientalis (L.) Link.) at higher elevations, but there are also pure Oriental beech  stands that cover large areas and are connected by corridors [4], [5], [6], [9]. Oriental beech forests, however, have been degraded, especially near villages and transport corridors [10]. Plus trees are very difficult to find in these areas. These circumstances can negatively affect natural regeneration practices. In this species, seedling establishment and growth rate depend on the seed tree, habitat conditions, and especially on the light factor [11]. Many studies on beech regeneration have been conducted in pure beech forests [12]. [13]. [14]. Studies on the regeneration of many plant species, especially beech, are usually correlated with light and weather conditions.

Ozel [14] stated that the group regeneration works in Bartin and Devrek in Turkey failed and that the areas that had lost the natural regeneration conditions needed regenerating by planting under the shelter of trees. Moreover, the seed is a very important material for outdoor plantation conditions, rehabilitation and conversion to high forest practices. Due to a lack of seed orchards for Oriental beech, the selected seed stands have been used in Turkey [15]. Generally, mast years occur every 3-5 years [12], [16], 5-6 years [4] for Oriental beech. However, Tosun [17] studied the seed yield of Oriental beech at Bolu (Turkey) and in the period of eleven years observed one mast year, two good years, three moderate years and five weak years. Suner [12] indicates that lower seed productivity exists in Oriental beech stands on calcareous bedrocks, whereas the seed year occurs more frequently and abundantly. Consequently, investigations on the seeds of this species have been on the increase because of rare mast years and germination problems caused by physiological dormancy [18], [19]. Moreover, Rezaei et al. [20] stated that oriental beechnuts are deeply dormant seeds that can germinate after 8-19 weeks of cold stratification. Besides, there were various studies on seed characteristics [21], [22], [23], [24], [25] and the storability of beechnuts [18], [26]. Thus, the aim of this study is to provide new information for seed technology of Oriental beech by examining the effect of tree age, stand exposures and elevations on seed characteristics.




Oriental beech belongs to the Fagaceae family. It is a shade-tolerant species [27]. Seedlings are tolerant to shade and grow under trees slowly, and therefore they exhibit growth with more speed in the juvenile age. The seedlings of the Fagus orientalis L. can grow in gravel soil and gravel-silt and are able to expand their own roots. The optimum pH of this species is 5.5 in the periods, and beech seedling that expand its roots, may be spherical. The density of seedlings has a direct impact on the form of the seedlings [3], [4], [6], [9].

Sampling Sites

The seeds of Oriental beech were collected in a natural forest in Kumluca - Bartın (41º30’5” - 41º20’27” N, 32º23’46” - 32º33’44” E) located in the northern part of Turkey (Figure 1). Oriental beech appears naturally from 400-1000 m a.s.l., especially in the Kumluca forest region [28]. Kumluca Oriental beech forests area was separated into two blocks on the basis of 600 m and 800 m elevations. In each block, 3 plots, in size of 100 m2, were taken according to age groups (40-59, 60-79 and 80-99 ages). In each plot, 3 trees with 150 m intervals were selected for having their seeds collected. The collected seeds were sampled, labelled and put in plastic bags. In addition, to determine the effect of the exposure factor, the seeds were obtained from 3 trees only from 60-79 ages group according to north, west, east and south exposures.


FIGURE 1. The locations of sampling sites at Kumluca-Bartin in Turkey


Studies under the Laboratory Conditions

The seeds, collected from these sites, were stored frozen until laboratory studies were initiated. In order to find out the amount of filled seeds, whole seeds were immersed in water and the ones floating on the surface of the water were separated. The filled seeds were chosen as 3×100 seed patterns for each sampled tree. In order to determine the weight of 100 seeds of Oriental beech, an average of three groups of 100 seeds was weighted. In the method for testing moisture content; three seeds group were separated and weighted for their wet and dry weights. The moisture contents (WA) were calculated [9] as follows:

WA= (A-E) • 100/A

where A is the wet weight and E is the dry weight.

The seeds were stratified in moisture-sand in a cool room at 4±0.5ºC for 90 days [9], [27].

Studies under the Greenhouse Conditions

The number of 3×100 seeds for each sampled tree sowed at ¼ sand+ ¼ clayey soil + ½ peat in the depth of 2-3 times of its size in the greenhouse in April. Following the sowing, the medium was thoroughly watered and kept damp all the time. Following the sowing date, the greenhouse temperature was kept at 25ºC and 85% humidity. The germination period was completed in 45 days.

Statistical Analysis

All experiments were based on completely random designs with three replicates, using 100 seeds in each replicate. The germination percentage was calculated as the proportion of the germinant of the total number of seeds. Arcsin (p)1/2 transformed germination data were subjected to analysis of variance using computer software package SPSS 9.0 and Duncan’s multiple range test was performed for significant effects.



The Impact of Elevation

The results of ANOVA for the effect of elevation factor on seed germination and moisture content showed significant differences at 0.001 confidence level (Table 1).

TABLE 1. Analyses of variance for elevation factor

The elevation factor was especially very effective on the seed germination of Oriental beech. As a matter of fact, while the seeds collected from 800 m elevation showed 72% germination, other seeds, collected from 600 m elevation, presented 57% germination. Also, the moisture contents were found to be 14% for 600 m and 16% for 800 m elevation (Figure 2).

FIGURE 2. The elevations factor on seed germination and moisture content

Saatcioglu [9] says that Oriental beech has fairly low moisture content (10.6-12.0%). In this study, the result obtained was very similar. It is informed that high germination as 80% was obtained from low moisture content [9]. However, the high germination ratio in this study was obtained from high moisture content. The germination percentage of 800 m elevation was higher than of 600 m elevation. The 800 m elevation degree has nearly optimum distribution elevation for Oriental beech in this ecologic region. According to Vahid et al., most of the seed production in Oriental beech takes place in 750 m above sea level [23].

In this study, 100 seed weight of 600 m and 800 m were 29.63 g and 27.63 g respectively. Oriental beech has heavy seeds and rarely has a mast year compared to some other forest trees. Saatcioglu and Urgenc [29] informed that the weight of 1000 seeds of Oriental beech was 273.1 g (215.3-316.7). These results are in accordance with the information by Saatcioglu and Urgenc [29], as well as by Gezer and Yucedag [19].

The Impact of Tree Age

According to the results of ANOVA, tree age factor had a significant effect on seed germination at 0.01 confidence level (Table 2). The highest and lowest germination percentages were determined as 76% and 54% on trees aged 60-79 and 40-59 respectively (Figure 3).

TABLE 2. Analysis of variance of tree ages factor

FIGURE 3. The effects of tree age’s factor on germination percentage

Farmer [30] informed that Fagaceae languish 2 or 3 decades before flowering. Older trees also may not reproduce if they are in the understory of stand. Therefore, the amount of seeds is first of all a function of the number, the age, the size and the physiological condition of the trees in the stand. Espahbody et al. [31] informed that more fertile seeds in terms of quality and quantity were obtained from middle aged trees. Although Fennessy [32] said that high quality fertile seeds increased from higher age to middle age, especially the beech and the oak tree, in this study, it was apparent that tree ages of beech merely affected the seed germination positively. On the other hand, Espahbodi et al. [31] confirmed that there was high germination on Sorbus seeds which were obtained from middle aged trees. Similar results to ours were also found on fir species by numerous researchers [33].

The Impact of Exposure

According to the results of ANOVA, the exposure factor had a significant effect on seed germinationat 0.001 confidence level, on moisture content at 0.01 confidence level and on 100 seed weight at 0.05 confidence level (Table 3).

TABLE 3. The result of ANOVA for exposure factor on seed of Oriental beech


The results of Duncan test - the variation coefficient, the average value, homogenous groups of the exposure factor on the germination percentage, 100 seed weight and the moisture content are presented in Table 4.

TABLE 4. Average value, variation coefficient (CV) and homogeny groups of exposure factor on seed characteristics of Oriental beech

The highest germination percentage of 82% was obtained from the northern exposure, while the lowest germination was 65% from the western exposure (Figure 4).

This result is in accordance with the result of the research of the Oriental beech in Mazandaran Forests. It is concluded that the highest amount of collected seeds is related to the northern exposure of all studied trees [23].

FIGURE 4. The effects of exposure factor on germination percentage

While the effects of elevation and the tree age factors on 100 seed weight had no significance, the effect of the exposure factor had showed significant noteworthy results. As a matter of fact, while the highest 100 seed weight was obtained from the southern exposure, the lowest 100 seed weight was obtained from the eastern exposure. Soltani [34] informed that there were positive relations between the seed weight and the time of overcoming seed dormancy in beechnuts. This information was given by various researchers [35], [36]. However, this does not mean that the highest germination percentage cannot be obtained by earlier overcoming of seed germination. Furthermore, in this study, the highest germination percentage was supplied by the average seed weight and by northern exposure. From the observation of the samplings, it was determined that the beech trees in the northern exposure were healthier than others. In addition, technical staff of the general directorate of forestry said that the cavity of beech wood in the northern exposure was smaller than in other exposures.



Since Oriental beech seedlings are produced by the generative propagation method, seeds should be harvested in the optimum distribution area of beech, which is 800 m a.s.l., from average ages (60-79 years old) and in the northern exposure for the highest germination percentage.


  1. SAGHEB-TALEBI K, MIRKAZEMI Z, AKHAVAN R, KARIMIDOOST A, MAGHSOUDLOO MK, MOGHADASI D 2011 Some structural characteristics in the Far East border of the distribution range of oriental beech (Fagus orientalis Lipsky) stands. In: Wagner S,  Fahlvik  N, Fischer H (eds) Proceedings of The 9th IUFRO International Beech Symposium organized by IUFRO working party 1.01.07 “Ecology and Silviculture of Beech”, Dresden/Göttingen, Germany, 12-17 September 2011. Institute for Silviculture and Forest Protection, Dresden, Germany, pp 61-63
  2. ERCANLI I, KAHRIMAN A, YAVUZ H 2014 Dynamic base-age invariant site index models based on generalized algebraic difference approach for mixed Scots pine (Pinus sylvestris ) and Oriental beech (Fagus orientalis Lipsky) stands. Turk J Agric For 38: 134-147. DOI: http://dx.doi.org/10.3906/tar-1212-67
  3. SAATCIOGLU F 1969 Si1vicuture I, Biological Fundamentals and Principles of Silviculture. Istanbul University, Faculty of Forestry, 138, Istanbul, Turkey, 323 p
  4. ATAY İ 1982 Natural Regeneration Methods II, Dogal Gençleştirme Yöntemleri II. Istanbul University, Faculty of Forestry, Pub. No: 320, Istanbul, Turkey, 160 p
  5. ATALAY İ 1992 Kayın (Fagus orientalis ) Ormanlarının Ekolojisi ve Tohum Transferi Yönünden Bölgelere Ayrılması. Orman Bakanlığı, Orman Ağaçları ve Tohumları Islah Araştırma Müdürlüğü, Yayın No: 5, Ankara, Turkey, 209 p
  6. ATA C 1995 Silviculture Techniques (in Turkish). Zonguldak Karaelmas University, Bartın Faculty of Forestry, No: 3-4, Bartın, Turkey, 449 p
  7. OZEL HB, ERTEKIN M 2011 Growth models in investigating oriental beech (Fagus orientalis ) juvenilities growth performance in the Western Black Sea in Turkey (Devrek-Akçasu Case Study). Romanian Biotechnological Letters 16 (1): 5850-5857
  8. YILMAZ M 2010 Is there a future for the isolated oriental beech (Fagus orientalis Lipsky) Forests in Southern Turkey? Acta Silv Lign Hung 6: 111-114
  9. SAATCIOGLU F 1971 Si1vicuture Techniques II.IstanbulUniversity, Faculty of Forestry,172, İstanbul, Turkey, 554 p
  10. ANONYMOUS 2006 Forest Ministry of Environment and Forestry, General Directorate of Forestry, Ankara, Turkey, 152 p
  11. KEIVAN S, SAGHEB-TALEBI K, SOMAYEH A 2011 Characteristics in oriental beech seedlings in pure beech Forests (Case Study: Iran-Mazandaran-Liresar Forestry Plan, Fifth District). World Applied Sciences Journal 12 (5): 591-599
  12. SUNER A 1978 Research on natural regeneration of pure Oriental beech stands at Düzce, Cide and Akkuş regions. Forestry Research Institute, Technical Bulletin, No: 107, Ankara, Turkey, 60 p
  13. NAKASHIZUKA T, NUMATA M 1982 Regeneration process of climax beech forests, I. Structure of a beech forest with the undergrowth of Sasa. Jpn J Ecol 32: 57-67
  14. OZEL HB 2007 Stand Structures and the Factors Affecting the Success of Group Natural Regeneration Practices in Bartın and Devrek Oriental beech (Fagus orientalis ) Forests. PhD thesis, Zonguldak Karaelmas University, Institute of Science, 292 p
  15. ANONYMOUS 2004 Working Report of 2003 and Working Programme of 2004. Forest Tree Seeds and Tree Breeding Research Directorate, Ankara, Turkey, 15 p
  16. ATAY İ 1990 Silviculture II - Silviculture Techniques. Istanbul University, Faculty of Forestry, Pub. No: 405, Istanbul, Turkey, 242 p
  17. TOSUN S 1992 Studies on the seed yield of Oriental beech (Fagus orientalis ) at Bolu. Forestry Research Institute, Technical Bulletin, No:232, Ankara, Turkey, 75 p
  18. YILMAZ M 2005 The researches on the physiology of oriental beech (Fagus orientalis ) seeds. PhD thesis, Istanbul University, Faculty of Forestry, Istanbul, Turkey, 170 p
  19. GEZER A, YUCEDAG C 2006 Forest tree seeds and techniques of seedling propagation from seed originated. Suleyman Demirel University, Faculty of Forestry, Pub. No: 56, Isparta, Turkey, 140 p
  20. REZAEI A, NASERY B, YAZDIAN F, MHEDAYATI MA 2011 A study of gibberalic acid effect on oriental beech (Fagus orientalis) nuts dormancy removal. In: Wagner S, Fahlvik  N, Fischer H (eds) Proceedings of The 9th IUFRO International Beech Symposium organized by IUFRO working party 1.01.07 "Ecology and Silviculture of Beech", Dresden/Göttingen, Germany, 12-17 September 2011. Institute for Silviculture and Forest Protection, Dresden, Germany, pp 61-63
  21. GEZER A 1983 Fundamentals of seed and seedling production of beech and spruce. Journal of Forestry Engineering 20: 11
  22. FALLERI E, MULLER C, LAROPPE E 2004 Effect of water stress on germination of beechnuts treated before and after storage. Can J Forest Res 34(6): 1204-1209. DOI: http://dx.doi.org/10.1139/x04-003
  23. VAHID E, REZA MM 2004 Investigation on quality and quantity of seed production of beech (Fagus orientalis Lipsky) in Mazandaran Forests. In: Sagheb-Talebi K, Madsen P, Terazawa K (eds) Improvement and Silviculture of Beech, Proceedings from the 7th International Beech Symposium, Tehran, Iran, 10-20 May 2004. Research Insitute of Forests and Rangelands, Tehran, Iran, pp 57-60
  24. TABARI M 2008 Germination and initial growth of Fagus orientalis seedling under different stand canopies. Journal of Applied Sciences 8 (9): 1776-1780
  25. YILMAZ M, DIRIK H 2008 Maturation of oriental beechnuts (Fagus orientalis Lipsky). Dendrobiology 60: 57-62
  26. YILMAZ M 2008 Three-year storage of oriental beechnuts (Fagus orientalis Lipsky). Eur J Forest Res 127: 441-445. DOI: http://dx.doi.org/10.1007/s10342-008-0227-5
  27. URGENC S 1998 Tree and Shrub Plants, Nursery and Propagation Technique. Istanbul University, Faculty of Forestry, 3395-3442, Istanbul, Turkey, 640 p
  28. ANONYMOUS 2003 Forest Management Plan of Kumluca Region. Bartın Forestry Enterpries, Bartın, Turkey
  29. SAATCIOGLU F, URGENC S 1960 Effects of cold-wet method on germination of oriental beech (Fagus orientalis ) seeds. Istanbul University, Journal of Faculty of Forestry 10: 2
  30. FARMER RE 1997 Seed Ecophysiology of Temperate and Boreal Zone Forest Trees. St Lucie Press, Florida, USA, 6 p
  31. ESPAHBODI K, HOSSEINI SM, NODOUSHAN HM, TABARI M, AKBARINIA M, SHOORAKI YD 2007 Tree age effects on seed germination in Sorbus torminalis. General and Applied Plant Physiology 33 (1-2): 107-119
  32. FENNESSY J 2002 The Collection, Storage, Treatment and Handling of Broadleaves Tree Seed. Reproductive Material URL: http://www.coford.ie/media/coford/content/publications/projectreports/cofordconnects/ConnectsNote4.pdf
  33. GATALAN G, PARADOS JA 1983 Genetics of the pinsapo. Ann For Sci 9: 185-208
  34. SOLTANI A 2003 Improvement of Seed Germination of Fagus orientalis Lipsky. PhD thesis (unpublished), University of Agricultural Sciences, Department of Silviculture, Umea, Sweden, 190 p
  35. MUMFORD PM 1990 Dormancy break in seeds of Impatiens glandulifera Royle. New Phytol 115 (1):171-175. DOI: http://dx.doi.org/10.1111/j.1469-8137.1990.tb00935.x
  36. SPRINGER TL, DEWALD CL, AIKEN E 2001 Seed germination and dormancy in eastern gamagrass. Crop Sci 41: 1906-1910. DOI: http://dx.doi.org/10.2135/cropsci2001.1906


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