GENETIC VARIABILITY OF WHEAT GERMPLASM REPRESENTED IN THE

SOUTH PANNONIAN REGION

Ivana RUKAVINA1, Sonja PETROVIĆ2, Tihomir ČUPIĆ3, Sonja MARIĆ2, Sunčica GUBERAC²

and Luka DRENJANČEVIĆ¹

1
Croatian Centre for Agriculture, Food and Rural Affairs, Institute for Seeds and Seedlings,

Osijek, Croatia
2
University of J.J.Strossmayer Osijek, Faculty of Agriculture, Osijek, Croatia
3
Agricultural Institute Osijek, Croatia

Corresponding author: Ivana Rukavina, Croatian Centre for Agriculture, Food and Rural Affairs,

Institute for Seed and Seedlings, Usorska 19, Brijest, 31000 Osijek, Croatia, Phone: +385 31 27

57 18; Fax: + 385 31 27 57 16; E-mail: [email protected]

Hereby I confirm that this scientific paper is original and not published before and also not sent

for publication to other journal.

1
 ABSTRACT

In this study, genetic variability was investigated among 50 winter wheat varieties (Triticum

aestivum L.) which are grown in parts of Croatia, Hungary, Serbia and Slovenia according to 22

morphological characteristics used for DUS (distinctness, uniformity and stability) testing. The

average Dice similarity coefficient was 0.371. The determined similarity coefficient was in range

0.083 – 0.776. A significant variability of 6.21% in the breeding programs according to period

was determined as well as significant variability of 3.10% between breeding programs. The

UPGMA clustering divided investigated varieties into four main clusters. Based on data analysis,

most distant varieties with best morphological characteristics were found which will provide

valuable resource of new parent's combinations in future breeding programs. This paper also

provided valuable assessment of morphological characteristics to define distinctness criteria in

the DUS examination of wheat.

Key words: wheat, cultivars, variability, DUS, similarity

Running title: VARIABILITY OF WHEAT FROM THE SOUTH PANNONIA

2
 INTRODUCTION

Common wheat (Triticum aestivum L.) is the most widespread cultivated plant species for

human consumption. In various forms wheat is used by more than one billion people in the world

and it is grown on more than 220 million hectares, with a total annual production of about 729

million tones (FAOSTAT, 2015). Continental Croatia is located in the southern part of the

Pannonian region with an average wheat production of about 645 000 tones, which is grown on

average at about 157 000 ha (CROATIAN BUREAE OF STATISTIC, 2015).

Genetic variability is one of the factors important for plant breeding. Gathering and

collecting germplasm resources and genetic variability represent foundation of any breeding

process (ALI et al., 2008; NEUMANN et al., 2011; NOVOSELSKAYA-DRAGOVICH et al.,

2011). In breeding process, the choice of suitable parents is extremely important to ensure a wide

genetic variability and thus allow selection of desirable genotypes from crossing (REIF et al.,

2005; BEDE and PETROVIĆ, 2006).

Testing and evaluation of varieties morphological differences is of a great importance for

breeding process because in classical breeding process determination of variability between

varieties is based on a large number of morphological characteristics. In the last years, the

numerous studies of genetic variation of morphological traits were conducted on a range of

different species (SMYKAL et al., 2008; ČUPIĆ et al., 2009; TASUNOVA et al., 2010; TUCAK

et al., 2011; LI et al., 2012; RUKAVINA et al., 2013; DENČIĆ et al., 2015). For most species

the first genetic maps were made using morphological markers and yet are still used today in

many cultivars. The morphological characteristics are also used to describe new varieties when

examining distinctness, uniformity and stability (DUS) in process of registration of new plant

varieties as well as in process of granting plant breeder's rights (JONES et al., 2003;

3
RUKAVINA et al, 2008). There are different opinions about the advantages and disadvantages of

using morphological characteristics. The main disadvantages of morphological characteristics are

the influence of environmental factors and developmental stage of the plant (WINTER and

KAHL, 1995), while the SMYKAL et al. (2008) reports about main reasons and benefits of using

morphological characteristics such as large number of characteristics and easier way of

monitoring and evaluation.

Analysis of morphological characteristics gave a clear insight into the existing diversity in

terms of breeding centers and a year of registration and directs towards the most diverse

genotypes that can be used as a parent lines for a new selection cycle. The goal of the present

study was to provide a clear assessment of the variability of wheat germplasm created in south

Pannonian region. This paper also provides valuable assessment of morphological characteristics

to define distinctness criteria in the DUS examination of wheat.

MATERIAL AND METHODS

Plant material and field trials

Research was conducted on 50 varieties of hexaploid wheat (Triticum aestivum ssp.

vulgare L.) from five breeding centers but with a broad genetic origin. The breeding centers are

located in continental Croatia. Varieties were selected based on market share and their

significance in the production. The 17 varieties were from Agricultural Institute Osijek (PIO), 15

from BC Institute for Breeding and Production of Field Crops Zagreb (BC), 12 from Agrigenetics

Ltd. (AG), 4 from Jost Seed-Research Ltd. Križevci (Jost) and 2 varieties from the Faculty of

Agriculture, University of Zagreb (AFZG) (Table 1).

Table 1

4
 Trials have also included standard example varieties for studied morphological

characteristics that are listed in the CPVO (Community Plant Variety Office) TP/003/4 Rev.2

(2011), protocol for wheat and their role was to clarify the expression of each of the observed

characteristics.

Field experiments were set up in field trials for DUS testing according to CPVO TP/003/4

Rev.2 (2011) on locations Osijek 45°32'N and 18°44'E (main location) and Klisa 45°46'N and

18°1'E (reserve location). At both locations trial was set up according to randomized block design

with 2 replications and plot size was 6.25 m². Each plot included 200 plants / m², a total of 1250

plants per basic plot. For this study, the evaluation of morphological characteristics was

conducted at both locations during the two growing seasons (2012/2013 and 2013/2014).

Additional trials for the characteristic - seasonal type were set up on location Klisa in two

years according to CPVO TP/003/4 Rev.2 (2011). For this purpose all investigated varieties were

sown in the spring time 2013 and 2014, out of frosts. Trials also included varieties Fidel and

Slejpner, examples for the specified characteristic. The regular maintenance and protection

measurements were conducted on trials.

Morphological characteristics

Variability of varieties was analyzed on the basis of 22 morphological characteristics:

plant growth habit, frequency of plants with recurved flag leaves, time of era emergence,

glaucosity of fleag leaf sheat, glaucosity of ear, glaucosity of neck, plant length, straw pith in

cross section, shape of ear in profile, density of ear, presence of awns or scurs, length of awns or

scurs at tip of ear, ear color, hairiness of convex surface of apical rachis segment, shoulder width

of lower glume, shoulder shape of lower glume, beak length of lower glume, extent of internal

hair of lower glume, grain color and seasonal type. Observations are done according to CPVO

TP/003/4 Rev.2 (2011) protocol for DUS testing of wheat (Table 2). During each vegetation year

5
seven morphological characteristics on 20 plants per plot were observed in field trials and the

characteristic seasonal type. At the harvest time, sample of 120 ears per plot were taken in order

to form samples of 20 ears for observation of 14 morphological characteristics: on the straw –

pith in cross section; on the ear - shape in profile, density, presence of awns or scurs, length of

awns or scurs at tip, color, hairiness of convex surface of apical rachis segment; on the lower

glume: shoulder width, shoulder shape, beak length, extent of internal hair and on grain

observation of color. In two growing seasons for each variety 1,120 plants were evaluated in the

field and 2,240 ears in the laboratory, which was in total, for 50 investigated varieties, 56,000

observations in the field and 112,000 observations in the laboratory. Investigated varieties in the

field trials were grouped according to the following characteristics recommended by CPVO

(2011): straw – pith in cross section (halfway between base of ear and the stem node below), ear

color, the presence of awns or scurs and seasonal type.

Table 2

Statistical analysis

Statistical analysis included the results from 21 morphological characteristics. Due to the

identical value for all varieties characteristic straw pith in cross section was excluded from the

statistical analysis because it would have no impact on the determination of similarity. Based on

the obtained notes the starting matrix was composed and used to calculate the Dice coefficient of

genetic similarity (FERGUSON and CARSON, 2007). Similarities were calculated using the

computer program NTSYS 2.2. (ROHLF, 2009). Similarity matrixes obtained from the

morphological data were used to create a dendrogram using Unweighted Pair Group of

Mathematics Average - UPGMA. The Mantel test was used for estimating the correlation

between dendrogram and original matrixes (MANTEL, 1967). Similarity matrixes of

morphological data were transformed to genetic distances for analysis of molecular variance -

6
AMOVA (EXCOFFIER et al., 1992) to determine the genotypic variance between and within the

assumed level structure of the tested varieties of winter wheat. Computer program Arlequin

ver.3.5. (EXCOFFIER & LISCHER, 2010) was used for calculation of molecular variance.

RESULTS

Based on the analysis of wheat varieties morphological characteristics data, classification

of investigated varieties was made according to the distribution of different expression states

(Supplement 1). The average Dice genetic similarity coefficient ( Sij ) was 0.371. Average

similarity value between breeding centers was ranged from 0.29 to 0.39. The lowest similarity

coefficient was between AG and AFZG (0.29), while even similarity was between AG and BC

(0.37), AFZG and BC (0.38), AFZG and Jost (0.37) and PIO and Bc (0.39). The highest

similarity within breeding center had Jost Seed-Research Ltd. (0.53) and lowest similarity had

Agricultural Faculty Zagreb (0,24). The highest similarity coefficient was determined between

varieties Cerera and Koleda (0.776), Aura and Zdenka (0.667), Bc Irena and Zdenka (0.655), Bc

Mira and Zdenka (0.625) and varieties Prima and Nina (0.615). The lowest similarity value was

0.083 between varieties Ilirija and Panonka (Supplement 2).

The UPGMA clustering (Figure 1) divided investigated varieties into four main clusters.

Cluster I was further divided into two sub clusters. First sub cluster consisted of AFZG Karla and

in second sub cluster were grouped Ilirija, Talia, Helia, Mihelca, Kalista, Matea, Anika, Ema,

Divana, Dea, Olimpija, Bc Lira, Cerera and Koleda. Cluster II comprised of Una, Barbara,

Žitarka and Super Žitarka, while varieties Lucija, Seka, Gabi, Srpanjka and Panonka were

grouped in cluster III. Cluster IV was further divided into four sub clusters. Variety Janica was

allocated into first sub cluster, Nina and Prima were grouped into second sub cluster. Third sub

cluster comprised of Adriana, Bc Elvira, Bc Mira, Alka, Banica, Bc Lidija, Nika, Zdenka and

7
Aura, while fourth sub cluster comprised of Mura, Atena, Bc Renata, Fiesta, Marija, Golubica,

Katarina, Aida, Felix, Zlata, Sana, Bc Irena, Bc Antea and Nova Žitarka. Correlation coefficient

between similarity matrix and dendrogram (0.68) was highly significant (P ˂ 0.001) after 1.000

permutations of Mantel test.

Figure 1.

AMOVA analysis determined the distribution of the total variance based on the

morphological characteristics data sets on levels of structure between breeding programs, within

the breeding program by period, and between varieties at the period of the program. Periods are

defined by year of release of varieties and the first period was until 2001 (including the year

2001), and the second period was after 2001. Largest share of variability (90.69%) was accounted

for the differences between the varieties at the period of the program (Table 3). There was also a

significant value of variability (6.21%) in the breeding programs according to period, and a

significant value of variability (3.10%) between breeding programs.

Table 3.

DISCUSSION

Evaluation based on morphological differences between wheat varieties has a great

importance for breeding because in the conventional breeding process determination of the

variability between varieties is carried out with the assessment of a large number of

morphological traits. Morphological characteristics are used when creating genetic maps, as well

as to control the initial population and separating generation (ŠATOVIĆ, 1999). Today is an

important application of morphological characters to describe new varieties when examining

distinctness, uniformity and stability (DUS) in the process of variety registration as well as to

plant variety protection (JONES et al., 2003; MARIĆ et al., 2004; RUKAVINA et al., 2008).

8
Many researchers reported about the need to combine morphological traits with biochemical and

molecular markers (ŠATOVIĆ, 1999; COOKE et al., 2003; COLLARD et al., 2005; BÖRNER,

2006).

This study determined relatively low average similarity value (0.371) among the

investigated varieties which indicates the great morphological variability of wheat germplasm

originally from continental Croatian as a south Pannonian region. Similar results were obtained

by MARIĆ et al. (2004), ALI et al. (2008), SALEM et al. (2008) and PETROVIĆ (2011.) who

found a relatively high distance between the tested varieties and UPGMA method showed the

presence of significant genetic variability. Contrary to these results, MACCAFERRI et al. (2007)

found the average genetic similarity value 0.73, and only a very distant line of durum wheat were

distinguished on the basis of phenotype that included morphological characteristics

recommended for DUS testing. The very low genetic similarity value (0.24) within breeding

center Agricultural Faculty Zagreb was the result of the work of several breeding programs from

which have been created investigated varieties.

Correlation coefficient of similarity matrix and dendrogram was 0.68 which indicate a

good relationship between the results of similarity matrix and morphological cluster analysis.

Genetic similarity of tested varieties based on morphological data showed clustering by the

variety type and in some clusters by the origin. It should be noted that the characteristic no.10

(straw pith in cross-section) had the same state of expression for all investigated varieties and as

such was excluded from the statistical analysis because it was determined that due to the

expression there was no effect on estimating of distinctness, which is in accordance with research

of JONES et al. (2003) and CABALLERO et al. (2010). Based on this knowledge appears that

there is a need for proposal to Technical Working Party for Agricultural Crops of UPOV

(International Union for the Protection of New Varieties of Plants) and Agricultural Expert Group

9
of CPVO (Community Plant Variety Office) to delete this characteristic no.10 as a grouping

characteristic and estimating distinctness in future revision of technical protocol for wheat DUS

testing. According to obtained dendrogram, in the first cluster were grouped varieties which had

present awns (Divana, Koleda, Cerera, Bc Lira, Olimpija, Dea, Ema, Anika, Ilirija, Talia, Matea,

Kalista, Mihelca and Helia), and in sub cluster was separately allocated variety with awns AFZG

Karla due to colored ear in stage of maturity. In this cluster sister lines Cerera and Koleda were

pointed out due to result of same crossing NE 7060 76Y335 / VG-19 (Zlatna dolina X Kavkaz)

and therefore it was expected for these varieties to have the highest similarity coefficient (0.776).

Barbara, Super Žitarka, Žitarka and Una were aligned in second cluster and classification of the

first three varieties can be associated with pedigree, because Žitarka is parental component of

varieties Barbara and Super Žitarka. Reason of grouping Una in this group can be associated with

the expression of the characteristics on the lower glume (shoulder width, shoulder shape, beak

shape and extent of internal hair) and grain color. All of these are high yielding varieties

belonging to the early to mid-early varieties, low to medium-high stem length, good quality and

resistance to lodging (BEDE, 1994; PETROVIĆ, 2011; DREZNER, 2012). In particular sub

cluster were grouped Seka, Lucija, Panonka, Srpanjka and Gabi belonging to the very early and

early varieties, with assessed very narrow shoulder of lower glume. It can be concluded that the

characteristics like date of ear emergence and shoulder width had an effect on the grouping of

these varieties besides pedigree since Srpanjka is one of the parents is in the varieties Seka,

Lucija and Gabi.

Fourth cluster included remaining varieties without awns, which were morphologically

heterogeneous with respect to the characteristics of the ear and lower glume. Variety Janica

allocated in a separate sub cluster, very close to the previous sub cluster, which can be associated

with a pedigree because one of its parents is Srpanjka, but unlike the previous sub cluster it had

10
strong expression of ear glaucosity as well as a different ear shape. Second sub cluster comprised

of sister lines Prima and Nina which were the result of same crossing Sana / Gala and these

varieties also had one of the highest similarity coefficient (0.615) which was expected. Grouping

of varieties in third sub cluster (Alka, Bc Mira, Bc Elvira, Adriana, Banica, Bc Lidija, Aura,

Zdenka and Nika), was under the influence of characteristic scurs length and medium-strong to

very strong hairiness of apical rachis segment. Fourth sub cluster comprised of morphologically

heterogeneous varieties, but it should be noted that variety Nova Žitarka was the only determined

as alternative type after two years of seasonal testing in field trials.

According to AMOVA analysis high variation between varieties per period of breeding

program was expected, since in the creation of varieties were used genetically different parents

and different selection criteria. Significant, but much lower proportion of the variability between

breeding programs can be linked to the fact that they relatively often use similar or partly shared

parental components during the process of creating a new varieties and in a very similar agro-

climate conditions. Similar reasons for higher variability within, rather than between the assumed

levels found ROUSSEL et al. (2004) for breeding centers in France, ROUSELL et al. (2005) for

European countries, and PETROVIĆ (2011) for Croatian and foreign wheat.

CONCLUSION

Our data analysis based on morphological characteristics showed similarity coefficient ranging

from 0.083 – 0.776 and it can be concluded that the use of morphological traits are of great

importance for estimating the criteria of distinctness in DUS testing of wheat as well as for

testing of genetic distances in wheat germplasm. On the basis of data analysis the farthest

varieties of best morphological characteristics were found and that will provide the successful

selection of new parent's combinations in future breeding programs.

11
 AUTHOR CONTRIBUTIONS

Creation and leading of field trials: Ivana Rukavina. Observation of qualitative characteristics and

measurments of quantitative characteristics in field and laboratory: Ivana Rukavina, Sonja

Petrović, Sunčica Guberac and Luka Drenjančević. Analysis of data: Tihomir Čupić assisted by

Sonja Marić. Wrote the manuscript: Ivana Rukavina, Sonja Petrović, Tihomir Čupić and Sonja

Marić.

ACKNOWLEDGEMENT

The authors are grateful to Institute for Seeds and Seedlings (Croatian Centre for Agriculture,

Food and Rural Affairs) which made it possible to carry out field trials.

Part of the presented work was supported by the Research Installation Project PHENOWHEAT

(No.2000) financed by Croatian Science Foundation.

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16
 GENETSKA VARIJABILNOST GERMPLAZME PŠENICE ZASTUPLJENE U JUŽNO

PANONSKOJ REGIJI

Ivana RUKAVINA1*, Sonja PETROVIĆ2, Tihomir ČUPIĆ3, Sonja MARIĆ2, Sunčica

GUBERAC² i Luka DRENJANČEVIĆ¹

Hrvatska
2
Sveučilište J.J.Strossmayer Osijek, Poljoprivredni fakultet, Osijek, Hrvatska
3
Poljoprivredni Institut Osijek, Hrvatska

IZVOD

U ovom radu je ispitivana genetska varijabilnost 50 sorti ozime pšenice (Triticum aestivum L.)

koje se uzgajaju u delovima Hrvatske, Mađarske, Srbije i Slovenije koristeći 22 morfološka

svojstva za DUS testiranje pšenice. Proučavana je germplazma iz pet oplemenjivačkih centara,

koja uključuje sorte registrovane od 1983. do 2010 godine. Prosečni Dice koeficijent sličnosti je

0.371. Koeficijent sličnosti je utvrđen u rasponu 0.083 – 0.776. Utvrđena je i značajna vrednost

udela varijabilnosti od 6.21% unutar oplemenjivačkih programa po periodu, kao i značajna

vrednost udela varijabilnosti od 3.10% između oplemenjivačkih programa. UPGMA grupisanje

razdvojilo je ispitivane sorte u četiri glavne grupe. Na osnovu urađenih analiza podataka utvrđene

su najudaljenije sorte najboljih morfoloških svojstava što će doprineti uspešnom izboru novih

roditeljskih kombinacija u budućim oplemenjivačkim programima.

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