Variation in Wheat ( Triticum spp.) Landraces from Different Altitudes of Three Regions of Turkey

-1

Variation in wheat (Triticum spp.) landraces from different altitudes of three

regions of Turkey

Alptekin Karagoz1 and Nusret Zencirci2,*1Genetic Resources Unit, Central Research Institute for Field Crops, P.O. Box 226, Ulus, Ankara-Turkey;2Department of Project Evaluation and Coordinator For IWWIP, CRIPC, P.O. Box 226, Ulus, Ankara,Turkey; *Author for correspondence (e-mail: [email protected])

Received 6 November 2003; accepted in revised form 20 September 2004

Key words: Landraces, Triticum spp., Turkey, Variation, Wheat

Abstract

Wheat landrace populations, collected from different altitudes of three regions of Turkey, were evaluatedfor variation within and among populations. Experimental material consisted of 380 accessions, from nineprovinces, in Central Anatolian Plateau and North Transitions. The material was grown at Haymana-Ankara Research Farm of Central Research Institute for Field Crops in 1999–2000 planting season, in athree-meter, two-row-plot trial with three bread and three durum wheat checks. Populations were evaluatedfor plant height, tillers number, spike length, grains per spike, awn length, thousand kernels weight (TKW),winter survival, and days to flowering. Observations and measurements were performed on five randomlyselected plants in each plots. Mean, coefficient of variation (CV), and range were computed for populationsfrom five altitude intervals of 0–399, 400–799, 800–1199, 1200–1599, and 1600–1999 m, and geographicalregions. Correlation, principal component (CPC) and cluster analyses were performed later. The highestvariation was recorded for awn length and the lowest for number of days to flowering. First three principalcomponents (PCs) accounted for 60.69% of the total variation. Cluster analysis for bread wheat, durumwheat, cultivated einkorn and cultivated emmer grouped the species meaningfully.

Introduction

Wheat is an important staple crop around theworld. Its importance has risen even more due tothe frequently experienced food shortages and itsrole in world trade. Increasing wheat productionto meet higher demands by growing population isstill a challenge in many countries. Higher pro-duction is only possible via higher yielding, betterquality, and disease resistant varieties. Successfulbreeding programs require wide variation. Turk-ish wheat landraces have been largely utilized tobreed new varieties through mass selection; Sivas

111–33, Sertak 52, Ak 702 (bread wheat) andKunduru 1149 (durum wheat) are some of them.And on the large variation existing in landraces(Porceddu and Scarascia-Mugnozza 1983; Blumet al. 1989; Vojdani and Meybodi 1993; Zencirciand Kun 1996) in comparison to the lower ge-netic variation in both current bread T. aestivum(Zencirci 1993; Zencirci 1998; Barbosa Neto andde Carvalho 2002) and durum wheat (T. durumDesf.) cultivars (Zencirci et al. 1994) for eco-nomic traits, have led breeders to search for newgenetic sources of variation (Wegrzyn et al.1996).

Genetic Resources and Crop Evolution (2005) 52: 775–785 � Springer 2005

DOI 10.1007/s10722-004-3556-3

Turkish wheat variation has always receivedgreat attention since the beginning of 20th century.Exploration and collection missions were mountedand the collected germplasm evaluated in differentcountries (Gokgol 1935, 1939; Harlan 1950;Zhukovsky et al. 1951).

The analyses of these material convinced Gok-gol (1935) that almost all wheat varieties existingin the world were present in Turkey and thatTurkish landraces provide an endless treasure tothe wheat breeders. Richness of wheat varieties ismainly due to the fact that diverse climatic andnatural conditions create a suitable environmentfor natural hybridization and recombination, thusgiving rise to new variation. In fact Gokgol (1939)identified almost 20,000 wheat types among theaccessions he examined with a number of thempeculiar of that origin.

Zhukovsky et al. (1951), considered Anatolia awarehouse where wheat varieties were assembledand filtered, before spreading to neighboringcountries.

Southeast Anatolia is known as primary genecenter of wheat diversification as well as the areaof first wheat domestication around 10,000 BP(Harlan 1981; Diamond 1997; Heun et al. 1997;Nesbit and Samuel 1998; Lev-Yadun et al. 2000).Mediterranean climate prevails in the region withcool and rainy winters, dry and hot summers. Thealtitude in Southeast Anatolia range between 350and 800 m, where both bread and durum wheatare widely grown. Central Anatolia plateau (900–1800 m a.s.l.) with its continental climate is themajor bread wheat growing area of Turkey. NorthTransition Zone (0–1800 m a.s.l) is characterizedby cool, high rainfall winters on coastal areas andcontinental climate in the inner part. Sharp cli-matic differences over short distances make pos-sible the accumulation of large variation in theregion. North Transition Zone variation is en-hanced by the cultivation of hulled wheatsT. monococcum L. (cultivated einkorn), T. dicocconSchrank (cultivated emmer), although their acre-age is steeply declining (Karagoz 1995).

Material and methods

Experimental material consisted of 380 wheatpopulations collected from farmers’ fields, village

threshing grounds and farmer stores of SoutheastAnatolia, Central Anatolia, and North Transi-tions, during 1986–1996 (Figure 1). Populationswere sampled to represent the variation present infarmers’ fields. Collection sites were selected fromremote villages, where landraces had still not beenreplaced by modem cultivars.

Accessions were planted on October 25th,1999 at the Haymana-Ankara Research Farm ofCentral Research Institute for Field Crops, in athree-meter, two-row-plots, unreplicated design(Delacy et al. 2000). Three bread (Bezostaya 1,Gun 91, and Ankara 98) and three durum wheatchecks (Kunduru 1149, Kiziltan 91, and AItin 40–98) varieties were added as checks.

Minimum temperatures of � 17.2 �C (January)and � 13.7 �C (March) and maximum tempera-tures of + 35.5 �C (August) and + 38.8 �C (July)were recorded. Total annual precipitation was332 mm.

The following traits were measured on fiverandomly taken plants, from each accessions(IBPGR 1985):

Plant height: The height of plant from groundlevel to end of spike excluding awns atmaturity (cm).

Spike length: The length of spike excludingawns at maturity (mm).

Awn length: The length of awns at maturity(mm).

Tillers number: Number of tillers counted afteruprooting.

Grains per spike: Number of grains counted onmain spike.

Winter survival: Observations on a 1–5 scale(1 = the lowest; 5 = the highest) just after winter.

Days to lowering: Number of days when 50% ofthe plants of each plot reached to flowering stage(January 1 = 1).

Thousand kernels weight (TKW): Averageweight (g) of 4 lots of 100 seeds, multiplied by 10.

Populations were grouped into three regions –Southeast Anatolia, Central Anatolia, and NorthTransitional Zone – and five altitude intervals – 0–399, 400–799, 800–1199, 1200–1599, and 1600–1999 m (Murphy and Witcombe 1981) – foranalyses.

Mean, coefficient of variation (CV), and rangewere computed for each accession, altitude, andregion. correlation, principal component (PCA)and cluster analysis were computed on averagevalues of accessions.

776

Results and discussion

Relatively large variation was observed for plantheight, tillers number, spike length, grains perspike, awn length, TKW, winter survival, and daysto flowering (Tables 1 and 2). Variation was largeenough to imply that populations could producesuitable genotypes to be utilized in breeding pro-grams (Figures 1 and 2). Values of single traits arethe following:

Plant height (cm)

Plant height means was equal to 99.11 cm with10.51% CV and 63.80–122.30 cm range. Dotlacilet al. (2000) considered a minimum 10% CV a signof wide diversity in wheat landraces and obsoletevarieties. Observed variation in plant height might,therefore be sufficient for an effective selection(Table 1, Figures 2 and 3). Bread wheats were thetallest followed by durums, einkorn and emmer(Table 2). Grouping samples into 400 m intervalsof altitude revealed that variation for this trait waslargest in samples from 800 to 1199 m intervals,and smallest in those from 1600 to 1999 m(Table 3). Samples from North Transitional Zonerevealed the largest variation with a CV of 11.50%for that trait, and samples from Central Anatoliathe smallest with a % CV of 9.42 (Table 4). Plantheight was positively correlated with tillers num-ber, spike length, winter survival (p ‡ 0.01), num-ber of grains per spike (p ‡ 0.05), and negativelywith awn length and days to flowering (p ‡ 0.01)(Table 5). Although under low moisture stress,plant height is positively correlated with grainyield (Duwayri and Nachit 1989), in cooler areasplant height has no effect on grain yield(Moghaddam et al. 1998) as it was the case in thisstudy.

Tillers number

Plants’ mean tillers number was equal to 3.85, witha % CV of 28.76 and range of 1.00–8.00 (Table 1,Figures 2 and 3). There has not been a significant

Figure 1. Number of wheat populations with provinces of

origin.

b

777

distinction between the wheat groups for tillersnumber (Table 2). Variation was largest in sam-ples from 400 to 799 m intervals of altitude andand smallest in those from 1600 to 1999 m.Accessions from Southeast showed the largestvariation with a CV of 29.89%, and those fromNorth Transition the smallest ones with a 25.69%CV (Table 4). Tillers number was correlated withplant height and spike length (p ‡ 0.01) (Table 5).

Spike length (mm)

Mean spike length was equal to 73.78 mm with arange of 37.00–111.60 mm and 21.09% CV

(Table 1, Figures 2 and 3). The longest spikes wereobserved in bread wheat, followed by durum, em-mer and einkorn Table 2). Spike length has a sig-nificant effect on yield (Omar et al. 1979;Moghaddam et al. 1998) and it’s high heritability(Satyavart et al. 2002), make this trait a valuableselection criteria for high grain yield. Variation waslargest in samples from 800 to 1199 m intervals ofaltitude and smallest in those from 0 to 399 m(Table 3). Samples from North Transitional Zoneshowed the largest variation with a CV of 25.85%,and those from Central Anatolia the smallest onewith 18.85% CV (Table 4). Spike length was sig-nificantly correlated with plant height, tillersnumber, grains per spike, winter survival

Table 2. Mean, minimum, maximum, SD and CV for plant height, tillers number, spike length, grains per spike, awn length, TKW,

winter survival, and days to flowering values for four different species.

Species Plant height Tillers number Spike length Grains/spike Awn length TKW Winter survival Days to flowering

T. aestivum Mean 100.47 3.95 78.82 30.52 44.45 35.94 2.88 149.10

Sx 10.42 1.07 12.95 6.73 31.60 4.06 0.74 2.72

Min 69.40 1.60 44.60 11.20 0.00 22.06 1.00 146.00

Max 122.30 7.60 111.60 62.40 135.60 45.43 5.00 160.00

CV 10.37 27.11 16.43 22.05 71.10 11.28 25.86 1.82

T. durum Mean 96.99 3.64 67.33 32.38 68.86 36.42 2.55 149.32

Sx 10.14 1.15 14.54 6.85 20.73 4.27 0.75 2.87

Min 68.30 1.80 38.20 15.60 0.00 25.73 1.00 147.00

Max 115.90 7.00 107.00 51.60 125.80 49.72 5.00 160.00

CV 10.45 31.65 21.59 21.14 30.11 11.72 29.31 1.92

T. dicoccon Mean 86.44 3.70 52.73 24.37 85.37 32.56 1.83 154.58

Sx 11.81 1.17 15.36 7.57 13.89 1.64 0.72 3.68

Min 63.80 1.00 40.00 15.40 58.40 29.46 1.00 150.00

Max 102.14 5.40 81.00 39.60 109.00 34.68 3.00 161.00

CV 13.66 31.56 29.13 31.05 16.27 5.05 39.15 2.38

T. monococcum Mean 92.83 4.25 51.03 26.35 70.90 32.26 2.63 156.13

Sx 9.82 1.07 15.68 9.17 14.09 3.71 0.52 6.51

Min 79.10 2.60 37.00 13.60 54.60 28.46 2.00 147.00

Max 109.44 5.80 81.00 36.60 94.80 38.83 3.00 164.00

CV 10.58 25.12 30.73 34.79 19.88 11.50 19.72 4.17

Table 1. Mean, CV, and range for plant height, tillers number, spike length, grains per spike, awn length, TKW, winter survival, and

days to flowering.

Traits Mean CV(%) Minimum Maximum

Plant height (cm) 99.11 10.51 63.80 122.30

Tillers number 3.85 28.76 1.00 8.00

Spike length (cm) 73.78 21.09 37.00 111.60

Grains per spike 30.93 22.18 11.00 63.00

Awn length (mm) 53.41 56.88 0.00 135.60

TKW (g) 35.81 11.44 22.06 49.72

Winter survival (1–5) 2.73 27.92 1.00 5.00

Number of days to flowering 149.58 2.19 146.00 164.00

778

(p ‡ 0.01), and negatively with awn length anddays to flowering (p ‡ 0.01) (Table 5).

Grains per spike

Mean values for grains per spike was equal to30.93 with 22.18% CV and range of 11.00–63.00.(Table 1, Figures 2 and 3). Durum wheat sampleshad the highest values, followed by bread wheat,emmer and einkorn Table 2). Variation for grainsper spike was largest in samples from 1600 to1999 m and smallest among those from 400 to799 m intervals (Table 3). Samples from NorthTransitional Zone had the largest variation with25.38 CV, and those from Southeast Anatolia thesmallest one with 18.97% CV (Table 4). Grainsper spike was significantly correlated with spikelength and awn length (p ‡ 0.01), negatively withwinter survival (p ‡ 0.01), with TKW and days toflowering (p ‡ 0.05) (Table 5). Since it affects

positively yield (Moghaddam et al. 1998) and ishighly heritable (Satyavart et al. 2002), grains perspike is used extensively as one of the majorselection criteria for higher grain yield.

Awn length (mm)

Mean awn length was equal to 53.41 mm with56.88% CV and range from 0 to 135 mm (Table 1,Figures 2 and 3). Average awn length was thehighest in emmer populations followed by einkorn(Table 2). Samples coming from 1600 to 1999 maltitude had the largest variation and those andfrom 0 to 399 mhad the smallest (Table 3). SamplesfromCentral Anatolia revealed the largest variationwith a 33.36% CV, and those from North Transi-tional Zone the smallest one with 24.80 CV %(Table 4). This large variation is mainly due tofarmers preference for awnless landraces e.g. Koseand its derivatives for their high bread making

Figure 2. Frequency distribution of populations for plant height, tillers number, spike length. grains per spike, awn length, TKW,

winter survival, and days to flowering.

779

quality in Sivas and Kayseri Provinces of CentralAnatolia and for awned landraces in Sanliurfa andAdiyaman in South East Anatolia (Table 4). Awnlength is significantly correlated with number ofdays to lowering andnumber of grains per spike, butnegativelywith plant height, spike length andwintersurvival (p ‡ 0.01) (Table 5).

Thousand kernels weight (g)

Average value was 35.81 g, CV 11.44% and rangewas 22.06–49.72 g (Table 1, Figures 2 and 3).

Figure 3. Coefficient of variation (%) for plant height, tillers

number, spike length, grains per spike, awn length, TKW,

winter survival, and days to flowering.

Table 3. Mean and CV far plant height, tillers number, spike length, grains per spike, awn length, TKW, winter survival, and days to

flowering computed on the basis of altitude groups.

Altitudes Plant height Tillers

number

Spike length Grains/spike Awn length TKW Winter

survival

Days to

flowering

Mean CV

(%)

Mean (%)

CV

Mean (%)

CV

Mean (%)

CV

Mean (%)

CV

Mean (%)

CV

Mean (%)

CV

Mean (%)

CV

0–399 99.9 10.11 3.73 27.66 73.17 6.98 31.57 21.88 52.26 20.53 32.83 8.70 2.67 38.73 151.00 2.18

400–799 95.32 9.68 3.64 30.12 73.97 21.62 33.23 18.89 62.56 31.54 37.77 10.00 2.69 24.76 147.91 1.37

800–1199 94.62 1l.88 3.75 28.75 64.51 23.49 30.05 23.50 76.65 23.21 33.13 14.16 2.21 40.19 152.35 2.91

1200–1599 101.85 9.88 4.02 27.81 76.10 20.32 30.53 21.40 43.47 76.61 34.78 9.55 2.80 24.10 150.14 2.08

1600–1999 104.77 8.76 3.99 27.26 75.33 16.82 26.83 27.36 31.94 93.77 37.41 9.93 3.22 24.45 148.92 1.59

Table 5. Correlation among populations far plant height, tillers number, spike length, grains per spike, awn length, TKW, winter

survival, and days to flowering.

Traits Plant height Tillers number Spike length Grains/spike Awn length TKW Winter survival Days to flowering

Trait # 1 2 3 4 5 6 7 8

2 0.290** – – – – –

3 0.288** 0.245** – – – – – –

4 0.121* 0.026 0.242** – – – – –

5 � 0.367** � 0.095 � 0.161 0.141 – – – –

6 0.078 � 0.016 0.089 � 0.116* 0.049 – – –

7 0.340 0.088 0.202 � 0.143** � 0.337 0.225** – –

8 � 0.144** 0.031 � 0.228 � 0.114* 0.189 � 0.433** � 0.263** –

Table 4. Mean and CV for plant height, tillers number, spike length, grains per spike, awn length, TKW, winter survival, and days to

flowering computed on the basis of geographical regions.

Regions Plant height Tillers

number

Spike length Grains/

spike

Awn length TKW Winter

survival

Days to

flowering

Mean (%)

CV

Mean (%)

CV

Mean (%)

CV

Mean (%)

CV

Mean (%)

CV

Mean (%)

CV

Mean (%)

CV

Mean (%)

CV

Southeast 95.46 9.49 3.68 29.89 74.53 21.12 33.24 18.97 62.74 30.06 37.89 9.97 2.66 25.42 147.88 1.38

Central Anatolia 102.81 9.42 3.98 28.62 75.80 18.85 29.88 22.33 41.64 80.10 35.57 9.82 2.90 24.63 149.64 1.82

North Transition Zone 93.86 11.50 3.77 25.69 65.08 25.85 29.75 25.38 74.93 24.80 32.29 12.81 2.27 39.04 152.96 2.88

780

Average values for this trait were close to eachother in different wheat groups while the maxi-mum values were obtained from durum wheat andminimum were from bread wheat and einkornTable 2). As for the altitudinal origins, the highestvariation was obtained from 800 to 1199 m andthe smallest was 0–400 m (Table 3). Samples fromNorth Transitional Zone showed the largest vari-ation with a CV of 12.81% and samples fromCentral Anatolia the smallest variation with9.82% CV (Table 4). Thousand kernel weight issignificantly correlated with winter survival, neg-atively with number of days to flowering(p ‡ 0.01), and number of grains per spike(p ‡ 0.05) (Table 5).

Winter survival (1–5)

Winter survival showed a mean score of 2.73 and27.91% CV and range between 1 and 5 (Table 1,Figures 2 and 3). Average winter survival score wasthe highest in bread wheat, followed by einkorn,durum and emmer. While full range of winter sur-vival score were observed in bread and durumwheats, the range was between 1 and 3 in einkornand 2–3 in emmer samples (Table 2). Variation waslargest in the samples from 800 to 1199 m and thesmallest in those from 1600 to 1999 m (Table 3).Samples from North Transitional Zone showed thelargest variation with a CV of 39.04%, and thosefromCentralAnatolia the smallest onewith a%CVof 24.63 (Table 4).Higher variation in samples fromNorth Transitional Zone might be due to sharpclimatic and altitudinal differences in short dis-tances, whereas lower variation in Central Anatoliawas a result of a relatively homogenous climaticcondition, with long-coldwinters.Winter survival issignificantly correlated with plant height, spikelength, TKW, and negatively with number of daysto flowering, awn length and grains per spike(p ‡ 0.01) (Table 5).

Number of days to flowering

Mean number of days to flowering was equal to149.58, with 2.19% CV and ranged between 146.00and 164.00 days (Table 1, Figures 2 and 3). Breadand durum wheat samples gave similar mean val-ues while emmer was few days earlier than ein-

korn. Range was the largest in einkornpopulations (Table 2). Variation was largest in thesamples from 800 to 1199 m and the smallest inthose from 400 to 799 m (Table 3). Samples fromNorth Transitional Zone revealed the largest var-iation with 2.88% CV and those samples fromSoutheast Anatolia showed the smallest one with1.38% CV (Table 4). Number of days to floweringwas positively correlated only with awn length,negatively with plant height, spike length, wintersurvival (p ‡ 0.01), and number of grains per spike(p ‡ 0.05) (Table 5). Lower variation for numberof days to flowering was not in agreement withdata by Spagnoletti Zeuli et al. (1983) and Zencirciand Kun (1996).

Principal component analysis

Though no clear guidelines exist to determine thesignificance of a coefficient, coefficients with ‡0.3 isregarded significant (Hair et al. 1987). Plant height(0.467), spike length (0.392) and winter survival(0.452) formed PC 1; tillers number (0.427), numberof grains per spike (0.413) and days to flowering(0.373) PC2; kernels number (0.620), awn length(0.451) and spike length (0.305) PC3. Cumulativevariance summed up to a total of 60.69%. PC1’sshare in that total was equal to 27.48%, PC217.38% and PC3 15.82% (Table 6).

Cluster analyses

Accessions from each species were grouped bycluster analysis. Accessions with similar valueswere considered as a single one. Their number wasreduced to 68 and 59, including checks, for breadand durum wheat, respectively, 10 for diploidhulled wheats and 14 for tetraploid hulled wheats,both with two checks. The four separatedendrograms are shown in Figures 4–7.

Bread wheat

Cluster analysis based on Euclidean SimilarityMatrix for bread wheat accessions formed fourmain clusters (Figure 4). The first two clusters (C1,C2) did not contain any of the check varieties.Both C1 and C2 had accessions from almost all

781

altitudes of all three geographic regions, exceptsamples from 0 to 399 m in C1 and from 1600 mand above altitudes in C2. Cluster 3 (C3) consistedof accessions from North Transitional Zone andcheck variety, Gun 91, an improved variety widelyadapted to North Transitional Zone. Cluster 4(C4), included accessions from Central Anatoliaand check variety, Bezostaya 1. Bezostaya 1 is awell adapted variety in the Central AnatolianPlateau.

Durum wheat

Cluster analysis based on Euclidean SimilarityMatrix for durum wheat formed five clusters

(C1–C5) (Figure 5). C1 consisted of five acces-sions and no check varieties, four of them werefrom 400 to 799 m range of Southeast Anatolia,and one from 1200 to 1599 m range of NorthTransitional Zone. C2 consisted of accessionsfrom Southeast Anatolia to North TransitionalZone with altitudial range of 400–1599 m and acheck. C3 consisted of accessions from 400 to1599 m range in both Southeast Anatolia andNorth Transitional Zones. C4 consisted ofaccessions from 0 to 1599 m range of all threegeographic regions. C5 consisted of two checks,Kiziltan 91 and Yilmaz 98, only. Kunduru 1149,the check originally selected from a landrace,may indicate its affinity with Turkish durumlandraces.

Figure 4. Clusters formed with bread wheat accessions from different altitudes of three geographic regions.

Table 6. Coefficients of the traits for the first three PCs and variation computed for by each of them.

Traits PC 1 PC 2 PC 3

Plant height 0.467 0.269 � 0.129

Tillers number 0.245 0.427 � 0.096

Spike length 0.392 0.292 0.305

Grains per spike 0.043 0.413 0.620

Awn length � 0.387 � 0.039 0.450

TKW 0.259 � 0.554 0.295

Winter survival 0.452 � 0.206 � 0.248

Number of days to flowering � 0.382 0.373 � 0.383

Variation (%) 27.48 17.38 15.82

782

Figure 5. Clusters formed with durum wheat accessions from different altitudes of three geographic regions.

Figure 7. Clusters formed with cultivated emmer accessions.

Figure 6. Clusters formed with cultivated einkorn

783

Cultivated einkorn and emmer wheat

Although the number of accessions of cultivatedeinkorn and emmer was small, cluster analysiswere still preformed and dendrograms drawn.Two clusters were formed for cultivated einkornand two for for tetraploid ones (Figures 6and 7).

Conclusions

Large scale trait evaluations may enhance utili-zation of plant genetic resources collections byincreasing genetic variability for economicallysignificant traits into wheat breeding programs.Regions of diversity are the most appropriateplaces to collect highly variable accessions. Awide genetic base, useful for breeding purposeswas found in this study for plant height, tillersnumber, spike length, grains per spike, awnlength, TKW, winter survival and days to flow-ering. Patterns of variation according to altitudeand geographical regions would ease the searchof traits of interest. In conclusion, Anatolianwheat landraces still hold a large capacity fordiversity.

References

Barbosa Neto J.F. and de Carvalho F.I.F. 2002. Genetic vari-

ability in common wheat germplasm based on COP. Genet.

Mol. Biol. 25: 211–215.

Blum A. Golan G., Mayer J., Sinmena B., Shpiler L. and Burra

J. 1989. The drought response of landraces of wheat from the

northern Negev Desert in Israel. Euphytica 43: 87–96.

Delacy I.H., Skowmand B. and Huerta J. 2000. Characteriza-

tion of Mexican landraces using agronomically useful attri-

butes. Genet. Resour. Crop Evol. 47: 87–96.

Diamond J. 1997. Location, location, location: the first farmers.

Science 278: 1243–1244.

Dotlacil L., Hermuth J., Stehno Z. and Manev M. 2000.

Diversity in European winter wheat landraces and obsolete

cultivars. Czech. J. Genet. Plant Breed. 16: 29–36.

Duwayri M. and Nachit M.N. 1989. Utilization of durum

wheat landraces to improve yield and yield stability in dry

areas. WIS 69: 5–8.

Gokgol M. 1935. Turkish Wheats, Vol. I. Ministry of Agri-

culture, Yesilkoy Seed Breeding Institute Publications. No: 7,

Devlet Press, Istanbul, Turkey (In Turkish), 436 pp.

Gokgol M. 1939. Turkish Wheats, Vol. II. Yesilkoy Seed

Breeding Institute Publications. No. 14, Tan Press, Istanbul,

Turkey (In Turkish), 955 pp.

Hair J.F.Jr., Anderson R.E. and Tatham R.L. 1987. Multi-

variate data analysis with readings. MacMillan Publ. Co.,

New York.

Harlan J.R. 1950. Collection of crop plants in Turkey. Agron.

J. 42: 258–259.

Harlan J.R. 1981. The early history of wheat: earliest traces to

the sack of Rome. In: Evans L.T. and Peacock W.J. (eds),

Wheat Science – Today and Tomorrow. Cambridge Press,

pp. 1–19.

Heun M., Schafer-Pregl R., Klawan D., Castagna R., Accerbi

M., Borghi B. and Salamini F. 1997. Site of Einkorn

domestication identified by DNA finger printing. Science 278:

1312–1314.

IBPGR 1985. Descriptors of wheat (revised). IBPGR Secre-

tariat, Rome.

Karagoz A. 1995. Agronomic practices and socio-economic

aspects of emmer and einkorn cultivation in Turkey. In:

Proceedings of the First International Workshop on Hulled

Wheats. 21–22 July 1995. IPGR1, Tuscany, Italy, pp. 172–

177.

Lev-Yadun S., Gopher A. and Abbo S. 2000. The cradle of

agriculture. Science 288: 1602–1603.

Moghaddam M., Ehdaie B. and Waines J.G. 1998. Genetic

variation for and interrelationships among agronomic traits

in landraces of BW from Southwestern Iran. J. Genet. Breed.

521: 73–81.

Murphy P.J. and Witcombe I.R. 1981. Variation in Himalayan

Barley and the concept of centers of diversity. In:AsherM.I.C.,

Ellis R.P., Hoyter A.M. andWhilehosueR.N.H. (eds.), Barley

Genetics IV. Edinburg Uni. Press, pp. 26–36.

Nesbit M. and Samuel D. 1998. Wheat domestication: ar-

caebotanical evidence. Science 279: 1433–1435.

Omar M.A., Shalaby E.E., Kassem A.A. and Abdelbary A.A.

1979. Variation, heritability, correlation, and predicted gains

from selection in wheat (T. aestivum). J. Agric. Res. 27(4):

159–163.

Porceddu E. and Scarascia-Mugnozza T. 1983. Genetic varia-

tion in durum wheat. In: Proceeding of 6th International

Wheat Genetics Symposium. Kyoto, Japan, pp. 241–252.

Satyavart A., Yadaya R.K. and Singh G.R. 2002. Variability

and heritability estimates in bread wheat. Environ. Ecol.

20: 548–550.

Spagnoletti Zeuli P.L., de Pace C., Benedetteli S., Lafiandra E.

and Porceddu E. 1983. Variation in durum wheat popula-

tions from different geographical origins: VI. Comparison

between advanced and primitive cultivars. In: Proceeding of

Breeding Methodologies in Durum Wheat and Triticale.

Tuscia Viterbo, Italy, pp. 17–28.

Vojdani P. and Meybodi M. 1993. Distribution and genetic

diversity of primitive bread wheats in Iran. In: Damania A.B.

(ed.), Biodiversity and Wheat Improvement. John Wiley and

Sons, pp. 44–48.

Wegrzyn S., Mazurkiewicz B. and Kuc I. 1996. Variation in

selected characters of varieties from the winter wheat col-

lections. Bulletin Institute of Hodowli, Krakow, Poland, pp.

181–182.

Zencirci N. 1993. Contributions of ancestral lines to modem bread

wheat cultivars. Doga: Turkish J. Agric. Forestry 17: 673–681.

Zencirci N., Aktan B. and Atli A. 1994. Genetic relationships of

Turkish durum wheat cultivars. Doga: Turkish J. Agric.

Forestry 18: 187–192.

784

Zencirci N. and Kun E. 1996. Variation in landraces of durum

wheat (T. turgidum L. conv. durum (Desf.) M.K. from

Turkey. Euphytica 92: 333–339.

Zencirci N. 1998. Genetic relationships of Turkish bread wheat

cultivars. Doga: Turkish J. Agric. Forestry 99(22): 333–340.

Zhukovsky P.M., Kipcak C., Nouruzhan H. and Turkistanli S.

1951. Ecological types and economic importance of Anatolian

wheat. Agricultural Structure of Turkey. Turkish Sugar Beet

Plants Publications No. 20: 158–214. (In Turkish), 887 pp.

785