**4. Discussion**

238 Genetic Diversity in Plants

and carpel scape width is significant at the 0.05 level, other all correlations are significant at

Fig. 3. Geographic positions of populations and results of cluster analysis

Morphological distance and grouping according to Penrose formula are shown that Table 3.

**Pop. No 1 2 3 4 5 6 7 8** 

the 0.01 level (Table 4).

**2** 1.0171

**3** 0.8408 1.0587

**4** 0.8715 1.2715 0.6238

**5** 1.1440 1.4252 1.5351 1.5285

**6 7.1725 5.7758 7.4618 7.2740 8.8849** 

**7 0.4673** 1.5966 1.3288 1.3045 1.6894 **8.4679** 

**8** 0.9812 1.4185 0.8796 0.8711 1.7620 **9.2148** 1.0019

Table 3. Morphological distance among populations according to Penrose Formula

0.9410 1.2547 0.7509 1.0335 1.6805 **7.2567** 0.6187 1.0099 1.8281 2.0630 2.8355 2.1007 2.7878 **4.6134** 1.6283 1.9982 0.6217 2.0264 1.5577 1.8446 2.3340 **8.0759** 0.5578 1.3256 **12 0.3029** 1.3819 1.1421 1.2455 1.5132 **8.0839** 0.7395 0.6935 1.5400 2.5653 0.8524 1.9061 1.9414 **10.3635** 1.3006 1.2208 0.5383 0.8897 1.0649 1.2452 1.0398 **5.2708** 0.9439 1.5747 0.9053 1.1575 1.1795 0.8243 1.1764 **7.5639** 0.5617 0.5652 1.3048 2.1365 1.3576 1.6116 1.1450 **9.9517** 1.1685 0.7097 **17 0.4078** 1.0211 1.0387 0.8375 1.4796 **7.0460** 0.5381 0.7332

According to results of the cluster analysis and variance analysis, Ilgaz1 population is very different to other populations. It could be because of its longitude and different ecological and genetical material condition. Results of the cluster analysis (Figure 2) were well accordance with morphological distances. For instance, morphological distances of Ilgaz1 were the highest than the others. Similarly population of Ilgaz1 is very different to other populations according to cluster analysis. These results could be used in preparation of gene map, seed transfer zones, determination of breeding populations, gene conservation areas, geographic variation and resulting of provenance trials of the species in short period. Preparation of forest gene maps and determination of seed transfer zones and geographical variation by morphological distance were also suggested by Yahyaoğlu et al 2001.

Genetic variation can be determined with morphological characters (Güney, 2009; Kulaç et. al., 2010; Şevik, 2010), isosymes analysis (Bilgen and Kaya, 2007; Turna, 2003) and DNA markers (Clark et. al., 2000; Goldstein, 1995). Many researchers use these methods for determination to genetic variation on *Abies* species; Messaoud et all. (2007) *Abies balsamea,*  Okada et all., (1973) *Abies sachalinensis,* Parker et all. (1981) *Abies balsamea* and *Abies lasiocarpa,* Kolotelo (1998) *Abies amabilis*, *Abies grandis* and *Abies lasiocarpa* e.c.

Shea (1990) reported that the variation among the populations is small (1,3%) but significant in *Abies Lasiocarpa*. Sorensen and Franklin (1977) reported that, year effect including interactions with places and trees in places made up an estimated 45 % of the variance in seed weight and 25% of the variance in cotyledon number. Among population genetic variance was much lower than within population variance, ranging from 6.6 to 6.8% for drought resistance traits to 7.8–14.0% for bud-break dates and a maximum of 10.0–17.9% for height growth traits to *Abies alba*. Therefore, genetic variance was predominantly within population (Sagnard et al, 2002).

The average genetic distance for all pair-wise comparisons between the ten populations of *Abies alba* in Italy was 0.014 (Parducci and Szmidt, 1997). 7,3% of the total genetic variation was due to differences among populations for gymnosperms (Hamrick et. al., 1992) and 10% for eigth Abies species (Shea and Furnier 2002). 13.3% of the total diversity is distributed among populations in *Abies alba* (Vendramin et al. 1999). Great variation was observed in the heterozygosity among the population studied and ranged from 0.010 *(A. pinsapo)* to 0.328 *(A. cephalonica).* The inter population genetic diversity was about 26% of the total genetic diversity. The average coefficient of gene differentiation (Gst) was 0.255, which means that approximately 26% of the total diversity of the Mediterranean firs exist among the populations. In particular, the geographical Area III (Turkey) has scored the highest value of Gst (25.8%), (Scaltsoyianne, 1999).

The proportion of genetic diversity among the populations of *Abies sachalinensis* is 1,5 % (El-Kassaby, 1992), populations of *Abies mariesii* is 2,6 % (Suyama et al. 1992) and populations of *Abies cephalonica* is 4.8% (Fady and Conkle 1993).

Conte (2004) reported that ANOVA analysis of *Abies nebrodensis* indicated that most of genetic variation resides within subsets (84%). More than 10% of the total genetic diversity was due to differences among populations of *Abies nebrodensis* (Vicario et, al., 1995).

Total percentage of genetic variation present in the population explained by interplot or among subpopulation differences is 0,35% of *Abies fraseri*. Thus, more than 99% of the genetic variation is due to within plot (i.e. tree to tree) variation (Diebel and Feret 1991). Most of the genetic diversity lies within populations to *Abies cephalonica* (Fady and Conkle 1993). Less than 10% of the total observed variation appeared among populations of *Abies cephalonica* (Hamrick, 1989) and the variation among the populations is 11% in *Abies alba*  (Vicario et al. 1995). Vendramin et al. (1999) reported that 13.3% of the total diversity is distributed among populations in *Abies alba.* On average, the genetic diversity among populations of *Abies* species has been found to be 6,3 % (Hamrick et al. 1992).

gene map, seed transfer zones, determination of breeding populations, gene conservation areas, geographic variation and resulting of provenance trials of the species in short period. Preparation of forest gene maps and determination of seed transfer zones and geographical

Genetic variation can be determined with morphological characters (Güney, 2009; Kulaç et. al., 2010; Şevik, 2010), isosymes analysis (Bilgen and Kaya, 2007; Turna, 2003) and DNA markers (Clark et. al., 2000; Goldstein, 1995). Many researchers use these methods for determination to genetic variation on *Abies* species; Messaoud et all. (2007) *Abies balsamea,*  Okada et all., (1973) *Abies sachalinensis,* Parker et all. (1981) *Abies balsamea* and *Abies* 

Shea (1990) reported that the variation among the populations is small (1,3%) but significant in *Abies Lasiocarpa*. Sorensen and Franklin (1977) reported that, year effect including interactions with places and trees in places made up an estimated 45 % of the variance in seed weight and 25% of the variance in cotyledon number. Among population genetic variance was much lower than within population variance, ranging from 6.6 to 6.8% for drought resistance traits to 7.8–14.0% for bud-break dates and a maximum of 10.0–17.9% for height growth traits to *Abies alba*. Therefore, genetic variance was predominantly within

The average genetic distance for all pair-wise comparisons between the ten populations of *Abies alba* in Italy was 0.014 (Parducci and Szmidt, 1997). 7,3% of the total genetic variation was due to differences among populations for gymnosperms (Hamrick et. al., 1992) and 10% for eigth Abies species (Shea and Furnier 2002). 13.3% of the total diversity is distributed among populations in *Abies alba* (Vendramin et al. 1999). Great variation was observed in the heterozygosity among the population studied and ranged from 0.010 *(A. pinsapo)* to 0.328 *(A. cephalonica).* The inter population genetic diversity was about 26% of the total genetic diversity. The average coefficient of gene differentiation (Gst) was 0.255, which means that approximately 26% of the total diversity of the Mediterranean firs exist among the populations. In particular, the geographical Area III (Turkey) has scored the highest

The proportion of genetic diversity among the populations of *Abies sachalinensis* is 1,5 % (El-Kassaby, 1992), populations of *Abies mariesii* is 2,6 % (Suyama et al. 1992) and populations of

Conte (2004) reported that ANOVA analysis of *Abies nebrodensis* indicated that most of genetic variation resides within subsets (84%). More than 10% of the total genetic diversity

Total percentage of genetic variation present in the population explained by interplot or among subpopulation differences is 0,35% of *Abies fraseri*. Thus, more than 99% of the genetic variation is due to within plot (i.e. tree to tree) variation (Diebel and Feret 1991). Most of the genetic diversity lies within populations to *Abies cephalonica* (Fady and Conkle 1993). Less than 10% of the total observed variation appeared among populations of *Abies cephalonica* (Hamrick, 1989) and the variation among the populations is 11% in *Abies alba*  (Vicario et al. 1995). Vendramin et al. (1999) reported that 13.3% of the total diversity is distributed among populations in *Abies alba.* On average, the genetic diversity among

was due to differences among populations of *Abies nebrodensis* (Vicario et, al., 1995).

populations of *Abies* species has been found to be 6,3 % (Hamrick et al. 1992).

variation by morphological distance were also suggested by Yahyaoğlu et al 2001.

*lasiocarpa,* Kolotelo (1998) *Abies amabilis*, *Abies grandis* and *Abies lasiocarpa* e.c.

population (Sagnard et al, 2002).

value of Gst (25.8%), (Scaltsoyianne, 1999).

*Abies cephalonica* is 4.8% (Fady and Conkle 1993).

The high within-population genetic diversity and low among-population differentiation observed in conifers have been attributed to common lifehistory traits, such as longevity and extensive gene flow (Hamrick et al., 1992; Streiff et al., 1998). The biogeographic history of a species should also contribute significantly to current patterns of genetic variation (Planter et al. 2000).

Despite the comparatively low levels of allozyme variation and the small genetic distances between populations, geographical differentiation among silver fir populations at different spatial scales could be demonstrated with markers (Konnert and Bergmann 1995). A large difference in cone legth, seed germination and seed weight was observed among the sites and among mother trees to *Abies sachalinensis* in Japan (Okada, 1973).

Contrary other Abies species, there are not enough study for Turkish fir. For this, it can be suggested that all populations, especially Ilgaz1 population, be considered for a gene conservation program. Also, future studies are necessary to provide deeper insights in to the subject. It may be concluded from the present study that studied characteristic were the important factors on morphological distance among populations. Ecological and geographical differentiation is important factors which influence the breeding and sampling strategies of tree crops. It is also essential to consider the relationship between population structure in natural and domesticated populations (Chalmers et al. 1992; Şevik, 2010; Şevik et al. 2011b). Results of this study could be taken into consideration in silvicultural purpose (afforestation, artificial regeneration) and breeding strategies (i.e. determination of breeding populations, gene conservation areas, seed transfer zones, seed sources and geographic variation, resulting of provenance trial; establishment of seed orchard) of this species.

Generally, our results show that large genetic diversity exist in Turkish fir to explain its great ecological plasticity and evolutionary. This results of study showed that the populations are not homogeneous with regard to the morphological characteristics. Populations consist of the trees having more or less different characteristics. The reason of the fact that the grouping and differences existed among the studied population in terms of the morphological characters may explain that there were different origins or varieties forming the Turkish fir stands. Variation in most of these characteristics appeared to be related altitude, divergent gene and genotype frequencies.

As is known, the morphological and physiological characteristics of forest trees are inherited. These features, with the growing effects of climate and environmental conditions can vary very little. As an example; needle length, the number of needles, cones, seed and leaf characteristics, branching characters as show some morphological features. In fact, many researchers in determining the genetic diversity of forest trees, one or a few of the uses of these characters. (Matziris, 1989; Cregg, 1994; Matziris, 1984; Komar, 2000; Fan ve Grossnickle, 1999; Lamhamedi, 2000; Matziris, 1997; John, 1948; Schmidtling et. al., 2005; Kathleen and Furnier 2002; Erkan, 2008; Bilir, 2002; Tylek ve Walczyk 2002, Güney et al., 2011, Kulaç et al. 2011a, 2011b; Turna and Güney, 2009; Turna, 2003, 2004). Seed size, parameters in terms of quality seeds is the most widely used classification also reflects the morphological diversity of values within and between populations. (Güney, 2009). Seeds in the trees, cones and cone elements, least affected by environmental conditions and thus genetic structure of the tree is considered the beginning of the elements that represent the most healthy way. Therefore especially in studies of genetic diversity of seeds, cones, and cones from the studies of the elements has a special place because it results is quite healthy (Turna et al., 2009).

Erkuloğlu (1993) reported that average weight of *Abies bornmulleriana* Mattf. Seeds from Bolu are 57,13 mg. Okada et al., (1973) *Abies sachalinensis* Masters in their study, in Japan, studied on 7 population and thousand grain weight of seed on the basis of population varied between 9.3 g to 12.3 g have identified. Also Skrzyszewska and Chlanda (2009) *Abies alba* Mill. thousand grain weight of seed on the basis of population, have found varied between 38.92 g and 53.27 grams. Edwards (1982), Fowells (1965)'to refer to *Abies lasiocarpa* var. *arizonica* (Merriam) Lemra. subalpine fir, compared to other types of seeds, the seeds of its much larger that represents about 70%. Kolotelo (1998) *Abies amabilis* seed weight (Dougl.) Forbes has changed between 25 mg and 55.6 mg and is the average of 34.5 mg, also *Abies grandis* Lindl. varied between 17.5 mg and 27.6 mg and average is 21.7 mg and *Abies lasiocarpa* (Hook.) Nutt. average of 7.2 mg to 18.5 mg and 12 mg of states that have changed. According to these results, the seeds of Uludag Fir, *Abies alba* Mill., *Abies amabilis* (Dougl.) Forbes, be said to be heavier than the seeds of *Abies lasiocarpa* (Hook.) Nutt. and *Abies grandis* Lindl. Also Sorensen and Franklin (1977) *Abies Procera* Rehd. state that the seed weight varied between 33 mg and 102.6 mg. In our study, the average seed weight ranged between 71.72 mg and 109.11 mg, the average was determined to be 81.58 mg. Gökmen (1970), indicates that *Abies nordmanniana* Mattf. seeds is 1 cm in length. According to these results, seeds of Uludag Fir with *Abies Procera* Rehd. seeds appear to be close to each other in weight. Franklin (1974) *Abies Procera* Rehd. carpel averaging 2.5 x 3 cm in size, seeds indicates that the average size of 12 x 6 mm. Macvean (2007) *Abies guatemalensis* Rehder indicates seed length about 8-10 mm and wings about 15 mm long.

Nowadays; because of its increasing economical value in market and decorative characteristic in landscape architecture, Turkish fir (*Abies nordmanniana* subsp. *bornmulleriana* Mattf.) is taking more importance. In addition to this being an endemic species of Turkey and widely preferred Noel tree in the world. Turkish fir is one of the most important trees to Christmas tree and for these there are very much studies on this species (Frampton and McKinley, 1999; Frampton et al. 2009; Talgø et. al., 2009; Newton et al., 2009; Hart et al., 2009; Langdren et al., 2008; Frampton and Işk, 2008; Talgø and Stensvand, 2008).

In this study, the genetic diversity of Turkish fir determined with respect to the some morphological characteristics. In this sense construction of genetic diversity, basic morphological characteristics, geographical variations and the morphological differences between tree species in the optimal and extreme distribution area of Turkish fir was determined.

Until now a few studies have been conducted about Turkish fir (Kaya et al., 2008; Şimşek, 1991; Velioğlu, 1999; Kaya and Raynal, 2000; Nielsen and Chastagner, 2005). But there is no comprehensive study to disclose the spatial distribution of Turkish fir and provide background information for future studies. In the near future; the studies done with the morphological characters about genetical variations are should also be analysed with DNA markers and isosymes analysis.

### **5. References**

Anonymous, 2006. Orman Varlğmz, T.C. Çevre ve Orman Bakanlğ Orman Genel Müdürlüğü, Orman İdaresi ve Planlama Dairesi Başkanlğ, Ankara, 160 s.

Basu, R.N. 1994. Seed viability. In *Seed Quality: Basic Mechanisms and Agricultural Implications*  (ed. A.S Basra), pp. 1-44. Food products press, New York.

Erkuloğlu (1993) reported that average weight of *Abies bornmulleriana* Mattf. Seeds from Bolu are 57,13 mg. Okada et al., (1973) *Abies sachalinensis* Masters in their study, in Japan, studied on 7 population and thousand grain weight of seed on the basis of population varied between 9.3 g to 12.3 g have identified. Also Skrzyszewska and Chlanda (2009) *Abies alba* Mill. thousand grain weight of seed on the basis of population, have found varied between 38.92 g and 53.27 grams. Edwards (1982), Fowells (1965)'to refer to *Abies lasiocarpa* var. *arizonica* (Merriam) Lemra. subalpine fir, compared to other types of seeds, the seeds of its much larger that represents about 70%. Kolotelo (1998) *Abies amabilis* seed weight (Dougl.) Forbes has changed between 25 mg and 55.6 mg and is the average of 34.5 mg, also *Abies grandis* Lindl. varied between 17.5 mg and 27.6 mg and average is 21.7 mg and *Abies lasiocarpa* (Hook.) Nutt. average of 7.2 mg to 18.5 mg and 12 mg of states that have changed. According to these results, the seeds of Uludag Fir, *Abies alba* Mill., *Abies amabilis* (Dougl.) Forbes, be said to be heavier than the seeds of *Abies lasiocarpa* (Hook.) Nutt. and *Abies grandis* Lindl. Also Sorensen and Franklin (1977) *Abies Procera* Rehd. state that the seed weight varied between 33 mg and 102.6 mg. In our study, the average seed weight ranged between 71.72 mg and 109.11 mg, the average was determined to be 81.58 mg. Gökmen (1970), indicates that *Abies nordmanniana* Mattf. seeds is 1 cm in length. According to these results, seeds of Uludag Fir with *Abies Procera* Rehd. seeds appear to be close to each other in weight. Franklin (1974) *Abies Procera* Rehd. carpel averaging 2.5 x 3 cm in size, seeds indicates that the average size of 12 x 6 mm. Macvean (2007) *Abies guatemalensis* Rehder

Nowadays; because of its increasing economical value in market and decorative characteristic in landscape architecture, Turkish fir (*Abies nordmanniana* subsp. *bornmulleriana* Mattf.) is taking more importance. In addition to this being an endemic species of Turkey and widely preferred Noel tree in the world. Turkish fir is one of the most important trees to Christmas tree and for these there are very much studies on this species (Frampton and McKinley, 1999; Frampton et al. 2009; Talgø et. al., 2009; Newton et al., 2009; Hart et al., 2009; Langdren et al., 2008; Frampton and Işk, 2008; Talgø and Stensvand, 2008). In this study, the genetic diversity of Turkish fir determined with respect to the some morphological characteristics. In this sense construction of genetic diversity, basic morphological characteristics, geographical variations and the morphological differences between tree species in the optimal and extreme distribution area of Turkish fir was

Until now a few studies have been conducted about Turkish fir (Kaya et al., 2008; Şimşek, 1991; Velioğlu, 1999; Kaya and Raynal, 2000; Nielsen and Chastagner, 2005). But there is no comprehensive study to disclose the spatial distribution of Turkish fir and provide background information for future studies. In the near future; the studies done with the morphological characters about genetical variations are should also be analysed with DNA

Anonymous, 2006. Orman Varlğmz, T.C. Çevre ve Orman Bakanlğ Orman Genel Müdürlüğü, Orman İdaresi ve Planlama Dairesi Başkanlğ, Ankara, 160 s. Basu, R.N. 1994. Seed viability. In *Seed Quality: Basic Mechanisms and Agricultural Implications* 

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