1 EVALUATION OF WHEAT GENOTYPES FOR SALINITY

2 TOLERANCE UNDER REAL SALINE CONDITIONS – IN SITU

3 Mirela Matković Stojšin1*, Sofija Petrović2, Borislav Banjac2, Velimir Mladenov2,
4 Veselinka Zečević3, Svetlana Roljević Nikolić1, Kristina Luković4
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Originalni naučni rad (Original Scientific Paper)
2 1
Matković Stojšin M, Roljević Nikolić S, Tamiš Research and Development Institute, Novoseljanski put
3 33, 26000 Pančevo, Serbia
4 2
Petrović S, Banjac B, Mladenov V, University of Novi Sad, Faculty of Agriculture, Dositej Obradović Sq.
5 8, 21000 Novi Sad, Serbia
6 3
Zečević V, Institute for Vegetable Crops, Krađorđeva 71, 11420 Smederevska Palanka, Serbia
7 4
Luković K, Center for Small Grains and Rural Development, Save Kovačevića 31, 34000 Kragujevac,
8 Serbia
9 Kontakt e-mail: matkovic.stojsin@institut-tamis.rs
 1 Summary

2 Salinity is a major abiotic stress factor that limits the productivity of crops, including

3 wheat, in many regions of the world. Therefore, developing wheat varieties that are

4 adapted to saline environments is a priority for global food security. In situ evaluation of

5 wheat genotypes can provide valuable information on the performance of different

6 genotypes under natural saline conditions and can help identify the most salt-tolerant

7 genotypes. To ensure an accurate evaluation of the performance of twenty-seven wheat

8 genotypes under different environments, the trial was conducted on two different soil

9 types (solonetz and chernozem) in two growing seasons. AMMI analysis shows that the

10 environmental factor had the largest share (55.15%) in the variation of grain yield, where

11 soil type had a dominant effect. Genotypes Renesansa, Harmonija, and Bankut 1205

12 achieved a high grain yield on both soil types. However, among the mentioned

13 genotypes, the genotype Harmonija showed the highest tolerance to salinity. A significant

14 proportion of the genotype and environment interaction (GEI; 25.89%) shows that there

15 is a change in the ranking of genotypes across environments. According to the AMMI 1

16 biplot, the genotypes Renesansa and Harmonija were distinguished by high grain yield

17 and high stability. The environment Chernozem 2015/2016 had the greatest contribution

18 to the GEI and the highest grain yield, while Solonetz 2017/2018 was characterized by

19 highest stability and the lowest grain yield. According to the AMMI 2 biplot, genotype

20 Harmonija achieved a high stability in unfavorable environmental conditions that

21 characterized environment Solonetz 2017/2018.

22 Key words: additive effects, multivariate effects, GEI, AMMI, salinity stress, stability

23
 1 Introduction

2 Wheat is one of the most widely grown and consumed cereals in the world, and it

3 provides around 20% of the calories and protein consumed by humans globally, making it

4 an important source of nutrition for millions of people (Shiferaw et al., 2013; Ernstein et

5 al., 2022). Given the widespread consumption of wheat-based foods around the world,

6 any disruptions to wheat production can have significant impacts on food availability,

7 prices, and nutrition. Therefore, ensuring the sustainability and resilience of wheat

8 production is critical for meeting the food needs of a growing global population (Lopes et

9 al., 2018). However, wheat production faces challenges such as extreme temperatures,

10 drought, soil salinity, and soil degradation (Shahid et al., 2018; Raimondo et al., 2020;

11 Saddiq et al., 2021). Soil salinity is one of the major abiotic stresses that negatively

12 impact plant growth and development (Dimitrijević et al., 2012; Farooq et al., 2022).

13 Different wheat genotypes can exhibit varying levels of tolerance to salinity stress, which

14 can affect their ability to produce acceptable grain yields under such conditions (Mansour

15 et al., 2020; Banjac et al., 2022). Evaluation of wheat genotypes under real field salinity

16 conditions—in situ—is crucial to identify salt-tolerant and sensitive ones, and this

17 information can be used to develop more effective breeding strategies for improved

18 wheat productivity in saline environments (El-Hendawy et al., 2017; Mansour et al.,

19 2020; Moustafa et al., 2021). Therefore, in order to identify wheat genotypes that are well

20 adapted to salinity stress conditions, it is necessary to conduct experiments in a variety of

21 environmental conditions. Many researchers used AMMI analysis with the aim of

22 assessing the stability of genotype performance across multiple environments (Petrović et

23 al., 2010; Dimitrijević et al., 2011; Mladenov et al., 2019; Verma and Singh, 2021;
 1 Banjac et al., 2022, Perišić et al., 2022; Ahakpaz et al., 2023). This method combines

2 ANOVA (Analysis of Variance) with principal component analysis (PCA) to separate

3 genotype and environment effects and their interaction (GEI) (Gauch and Zobel, 1996).

4 In the context of salinity tolerance in wheat, AMMI analysis can be used to evaluate the

5 performance of wheat genotypes under salinity stress and identify those that are more

6 tolerant (Petrović et al., 2010; Pour-Aboughadareh et al., 2021; Banjac et al., 2022;

7 Kumar et al., 2022; Farokhzadeh et al., 2022). This information can be used to develop

8 more effective breeding strategies for improving salinity tolerance in wheat, leading to

9 improved productivity and sustainability in saline environments.

10 The aim of this research is to: (I) establish the share of factors of genotype, environment,

11 and their interaction in the total variation of grain yield; (II) evaluate the stability

12 performance of the wider wheat germplasm across different environments; (III) identify

13 genotypes that exhibit satisfactory grain yield and high stability; and (IV) single out

14 genotypes that exhibit specific adaptability to salinity stress conditions.

15 Material and Method

16 Plant material and Experimental Design

17 Twenty-seven divergent wheat genotypes were used as material for the research,

18 including: two local populations (Banatka and Grbljanka), one old Hungarian variety

19 (Bankut 1205), 20 old and newly varieties created at the Center for Small Grains in

20 Kragujevac (Šumadija, Kosmajka, Gružanka, Morava, Zastava, KG-56, Orašanka, KG-

21 58, KG-78, Lepenica, Oplenka, Ljubičevka, Srbijanka, Šumadinka, Aleksandra, Perfekta,

22 Harmonija, Rujna, and Premija) and 4 varieties (Jugoslavija, NSR-5, Renesansa, and

23 Pesma) released by the Institute for Field and Vegetable Crops in Novi Sad.
 1 A trifactorial wheat trial was established, according to a completely randomized block

2 design, in three repetitions on two localities: Rimski Šančevi, near Novi Sad (Bačka) and

3 Kumane (Banat), during two growing seasons (2015/2016 and 2017/2018). Sowing was

4 carried out mechanically with a distance of 10 cm between rows, where the basic plot

5 size was 2 m2. During experimental research, the usual agrotechnics for wheat production

6 were applied. In both growing seasons, harvesting was done in the phenophase of full

7 maturity, when grain moisture dropped below 14%.

8 Soil conditions

9 Solonetz soil type is characterized by a high content of sodium and clay, which results in

10 a distinctive structure and chemical properties. The high sodium content of solonetz soils

11 can cause soil particles to bind tightly together, resulting in a hard, dense soil structure

12 that can be difficult for plant roots to penetrate. This can lead to reduced water and

13 nutrient uptake as well as decreased plant growth and yield (Belić et al., 2012). On the

14 other side, chernozem is a highly fertile and productive soil type that is well-suited for

15 agriculture. Chernozem soils are rich in organic matter, nutrients, and minerals, which

16 makes them highly productive and suitable for agriculture. They are formed from the

17 accumulation of organic matter over long periods of time, which creates a thick, black

18 layer of topsoil. Chernozem soils also have a unique physical structure, with a crumbly

19 texture that allows for good drainage and aeration (Hadžić et al., 2002).

20 Meteorologycal conditions

21 Meteorological data were obtained from the Republic Hydrometeorological Service of

22 Serbia's website (https://www.hidmet.gov.rs). The winter of 2015/2016 in the analyzed

23 localities was relatively mild, with temperatures above the long-term average. February,
 1 March, and April were characterized by warm weather, with temperatures and the sum of

2 precipitation around the long-term average, which had a positive effect on the growth and

3 development of wheat. The very high amount of rainfall in June allowed for a good

4 filling of the grain. However, in this period there was a wheat lodging that was

5 particularly related to old varieties and local landraces, which are characterized by a

6 higher stem height. The October, November, and December 2017/2018 growing seasons

7 were marked by higher temperatures compared to the multi-year average, with

8 significantly lower amounts of precipitation than average in both localities. February and

9 the first decade of March were characterized by significantly lower temperatures than the

10 multi-year average and a formed snow cover. At the end of March, there was a rise in

11 temperature, which favored the accelerated development of wheat. Very high mean

12 temperatures in April and May contributed to the drying of the surface layer of the soil

13 and a water deficit for the plants. June was characterized by temperatures within the

14 average range, with a higher amount of precipitation in the Rimski Šančevi locality. Dry

15 weather and high temperatures in the second half of the month accelerated the ripening of

16 wheat in both localities. In general, the 2017/2018 growing season was characterized by

17 lower rainfall and significantly higher average temperatures in April and May compared

18 to the 2015/2016 growing season in both localities (Figure 1).

19 Figure 1. Meteorologycal conditions during the experiment

20 Grafikon 1. Meteorološki usčovi tokom izvođenja eksperimenta

21 Statictical analysis

22 In order to analyze the influence of the analyzed factors and their interaction on the

23 variation of grain yield, an AMMI (Additive Main Effects and Multiplicative Interaction)
 1 analysis was carried out using the GenStat, Trial Version 18.1.0.17005

2 (https://www.vsni.co.uk). AMMI analysis combines elements of both ANOVA (Analysis

3 of Variance) and PCA (Principal Component Analysis) to decompose the genotype by

4 environment interaction (GEI) into additive main effects and multiplicative interaction

5 effects (Gauch and Zobel, 1996). The obtained PCA scores (IPCA1 and IPCA2) were used

6 to generate AMMI1 (IPCA1 vs. grain yield) and AMMI2 (IPCA1 vs. IPCA2) biplots,

7 which are useful graphical representations of the genotype and environment effects.

8 Results and Discussion

9 The AMMI model is considered an efficient method for analyzing GEI data in

10 agricultural research because it allows for the unique separation of main and interaction

11 effects, which is essential for identifying stable genotypes and making informed breeding

12 decisions (Gauch, 2006). The AMMI analysis of grain yield shows that there is a highly

13 significant (p<0.01) influence of additive and multivariate sources of variation. When

14 considering additive effects, the environment contributed the most to grain yield variation

15 (55.15%), while the genetic factor's part was significantly lower (11.16%) (Table 2).

16 Also, a greater share of environmental factors on wheat grain yield was established by

17 Verma and Singh (2021), and Pour-Aboughadareh et al. (2021). The analysis of

18 environmental factors showed that the soil type had a greater influence on grain yield

19 than the factor of vegetation season (Figure 2). Therefore, conditions of soil salinity

20 stress significantly reduced grain yield. This is in accordance with results established by

21 Dimitrijević et al. (2012), who stated that the reduction in grain yield was up to 40% in

22 alkaline soil. Also, Nadeem et al. (2020), Mansour et al. (2020), and Elfnah et al. (2023)

23 found that growing wheat on treatments with an applied high salt concentration
 1 significantly reduces grain yield, compared to the grain yield achieved on control and

2 treatments with a low salt concentration.

3 The least decrease in grain yield under the influence of salinity, was found in the

4 Šumadija and Rujna genotypes. However, although the most tolerant, the mentioned

5 genotypes achieved very low grain yields on both soil types. However, these genotypes

6 should not be immediately rejected as genetic material, but modern laboratory analyses

7 should be used to check whether they possess genes for salinity tolerance and whether

8 they can serve as a genetic resource in breeding for salinity tolerance. Genotypes

9 Renesansa, Harmonija, and Bankut 1205 achieved a high grain yield on both soil types

10 and are considered desirable genotypes for cultivation, both in favorable and unfavorable

11 environmental conditions. However, among the mentioned genotypes, Harmonija showed

12 the highest tolerance to salinity, i.e., the least decrease in grain yield under the influence

13 of stress factors, which is why it is considered particularly adapted to unfavorable salinity

14 conditions. On the other hand, the Zastava and NSR-5 genotypes, which had a large

15 difference in grain yield under the influence of salinity, achieved an above-average yield

16 on chernozem. Therefore, the mentioned genotypes are suitable for cultivation under

17 favorable environmental conditions, which will allow them to reach their full genetic

18 potential. When it comes to the factor of the growing season, genotypes Šumadija,

19 Premija, Gružanka, Harmonija, NSR-5, and Oplenka showed the least variation in grain

20 yield. Among the mentioned genotypes, the genotype Harmonija stands out as the most

21 productive. The highest difference in grain yield between growing seasons was recorded

22 in the local landraces Banatka and Grbljanka and in the old variety Bankut 1205, which

23 can be successfully grown under favorable environmental conditions (Figure 2).

 1 Figure 2. Grain yield variation under the influence of soil type (a) and vegetation season

2 (b) in the analyzed wheat genotypes

3 Grafikon 2. Varijacija prinosa zrna uslovljena tipom zemljišta (a) i vegetacionom

4 sezonom (b) kod analiziranih genotipova pšenice

5 The share of GEI in the phenotypic expression of grain yield was significant (25.89%),

6 which means that there was a change in the ranking of genotypes across environments

7 due to the large differences in the analyzed environments. Similar results were

8 established by Mohammadi et al. (2018), Wardof et al. (2019), and Verma and Singh

9 (2021). The interaction is decomposed into its interaction components, where the first

10 two main components (IPCA1 and IPCA2) explain 90.27% of the interaction (Table 1).

11 Table 1. AMMI-ANOVA for grain yield in different wheat genotypes grown on solonetz

12 and chernozem

13 Tabela 1. AMMI-ANOVA prinosa zrna različitih genotipova pšenice gajenih na

14 solonjecu i černozemu

15 The AMMI1 biplot (IPCA1 vs. grain yield) shows that the influence of additive and

16 multivariate variation on grain yield expression is almost equal (Figure 3). Genotypes or

17 environments that are plotted on the same vertical line of the AMMI 1 biplot do not differ

18 in yields, as they have similar additive effects. Similarly, genotypes or environments that

19 are positioned on the same horizontal line have similar interaction patterns and do not

20 differ in the multivariate part of the variation (Gupta et al., 2022). The environment

21 Chernozem 2015/2016, where the highest grain yield was achieved, had the greatest

22 contribution to the GEI. Agro-ecological environments that are characterized by less

23 favorable soil (solonetz) or less favorable meteorological conditions (Chernozem

 1 2017/2018) are located below the abscissa, where the environments of Solonetz

2 2015/2016 and Chernozem 2017/2018 differ from each other in the additive but not in the

3 multivariate part of the variation. The highest stability, but also the lowest grain yield,

4 was found in the environment of Solonetz 2017/2018. In this case, the high stability

5 resulted from the genotypes reduced capacity to express their genetic potential for grain

6 yield in unfavorable environmental conditions. The genotypes NSR-5, Srbijanka,

7 Lepenica, Ljubičevka, Šumadinka, KG-56, Zastava, Gružanka, Morava, KG-58,

8 Orašanka, Renesansa, and Harmonija showed high stability, i.e., low values of the first

9 interaction component. A genotype with both a high yield and a high degree of stability is

10 considered more desirable, as it has the potential to consistently perform well across

11 different environments (Kumar et al., 2022; Ahakpaz et al., 2023). Genotypes Harmonija,

12 Orašanka, Renesansa, KG-58, and Bankut 1205, in addition to their high stability, are

13 also characterized by a high average grain yield. Regardless of the high stability, the

14 genotype Kosmajka was characterized by low grain yield values in both localities.

15 Moderate stability was established in the genotypes Jugoslavija, Pesma, Banatka, KG-78,

16 Premija, and Šumadija, among which the genotype Jugoslavija stood out due to its high

17 grain yield. The genotype Rujna, in addition to the low value of the grain yield, was

18 distinguished by its pronounced instability (Figure 3).

19 Figure 3. AMMI1 biplot (IPCA1 vs. grain yield) for assessing the stability of wheat

20 genotypes grown in different agroecological environments

21 Grafikon 3. AMMI1 biplot (IPCA1 vs. prinos zrna) za procenu stabilnosti genotipova

22 pšenice gajenih u različitim sredinama

 1 By creating an AMMI2 biplot, an additional 28.13% of the interaction was explained

2 (Figure 4). In considering the fact that the AMMI2 biplot accounts for the majority of the

3 interaction, Khan et al. (2020) find that it is superior at estimating the complex GEI

4 pattern. Environments and genotypes that are positioned near the origin have less

5 influence, whereas those that are distant from the origin have a stronger impact on GEI

6 (Yan et al., 1998). In this study, genotypes that had low values of interaction components,

7 i.e., high grain yield stability across different environments, are: KG-58, Renesansa, KG-

8 56, Srbijanka, Lepenica, Ljubičevka, Šumadinka, Kosmajka, Grbljanka, NSR-5, and

9 Orašanka. The environments Chernozem 2017/2018 and Solonetz 2015/2016 had the

10 smallest contribution to GEI. The mentioned environments did not differ from each other

11 in the multivariate part of the variation; their vectors have the same orientation and give

12 the same ranking of genotypes. On the other hand, the environments Chernozem

13 2015/2016 and Solonetz 2017/2018, positioned in different quadrants of the biplot, had a

14 high effect on GEI. The importance of the growing season in achieving genotype stability

15 is reflected in the fact that the chernozem soil type in different growing seasons showed a

16 drastic difference in the multivariate part of the variation.

17 Genotypes with a higher IPCA score are more adapted to certain environments but less

18 consistent across different environments (Khan et al., 2020; Farokhazadeh et al., 2022;

19 Ahakpaz et al., 2023). In this research, the genotypes Rujna, Šumadija, Premija,

20 Harmonija, Pesma, Morava, Perfekta, and Aleksandra had high values of one or both

21 interaction components. However, some of the mentioned genotypes had positive

22 interactions with certain environments. The genotype Harmonija achieved a specific

23 adaptability to unfavorable environmental conditions that characterized Solonetz

 1 2017/2018 (Figure 4). Also, this genotype achieved the highest grain yield in saline soil

2 conditions in both analyzed growing seasons (Figure 2). Genotypes Aleksandra, Morava,

3 and Perfekta expressed stable reactions in the Solonetz 2015/2016 environment. The

4 genotype Pesma, together with the old variety Bankut 1205 and local landrace Banatka,

5 expressed specific stability in favorable conditions of the environment Chernozem

6 2015/2016 (Figure 4).

7 Figure 4. AMMI2 biplot (IPCA1 vs. IPCA2) for assessing the stability of wheat genotypes

8 grown in different agroecological environments

9 Grafikon 4. AMMI2 biplot (IPCA1 vs. IPCA2) za procenu stabilnosti genotipova pšenice

10 gajenih u različitim sredinama

11 Conclusion

12 The AMMI model can be a valuable tool for crop breeding and agricultural research,

13 helping to identify the best genotypes for specific environments. The results showed

14 significant influences of genotype, environment (soil type and vegetation season), as well

15 as GEI on the phenotypic variation of wheat grain yield. Regardless of low grain yields,

16 genotypes Rujna and Šumadija did not react to salinity stress conditions by reducing

17 grain yield. Genotypes Renesansa, Harmonija, and Bankut 1205 achieved high grain

18 yields on both types of soil, where the smallest decrease in stress conditions was recorded

19 in the genotype Harmonija. The favorable conditions of the Chernozem 2015/2016

20 environment allowed the genotypes to express their genetic potential, which resulted in

21 high yield instability in the given environment. On the other hand, the highest stability

22 and the lowest grain yield were found in the environment of Solonetz 2017/2018, where

23 genotypes did not have the capacity to express their genetic potential for grain yield.
 1 Genotypes Renesansa, Harmonija, KG‒58 and Orašanka expressed high above-ground

2 grain yield and stability and were considered suitable for growing across a range of

3 environments. According to the AMMI2 biplot, genotype Harmonija performed

4 specifically well under stressful conditions in Solonetz in 2017/2018 vegetation season,

5 and therefore it can be rated as a salinity-tolerant genotype.

6 Acknowledgments

7 This research was supported by the Ministry of Science, Technological Development,

8 and Innovation of the Republic of Serbia, grant numbers: 451-03-47/2023-01/200054;

9 451-03-47/2023-01/200117; and 451-03-47/2023-01/200216.

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18 Shiferaw B, Smale M, Braun H, Duveiller E, Reynolds MP, Muricho G (2013): Crops

19 that feed the world 10. Past successes and future challenges to the role played by

20 wheat in global food security. Food Sci. 5: 291–317. 

21 Verma A, Singh G (2021): Stability, adaptability analysis of wheat genotypes by AMMI

22 with blup for restricted irrigated multi location trials in peninsular zone of India.

23 Agric. Sci. 12: 198-212.

 1 Wardofa G, Mohammed H, Asnake D, Alemu T (2019): Genotype × environment

2 interaction and yield stability of bread wheat genotypes in central Ethiopia. J. Plant

3 Breed. Gen. Vol 7 (2): 87-94.

4 Yan W, Hunt LA (1998): Genotype by environment interaction and crop yield. Plant

5 Breed. Rev. 16: 135-178.

6
7 PROCENA TOLERANTNOSTI NA SALINITET KOD GENOTIPOVA PŠENICE

8 U REALNIM USLOVIMA SALINITETA – IN SITU

9 Zaslanjenost je glavni faktor abiotičkog stresa koji ograničava produktivnost useva,

10 uključujući pšenicu, u mnogim regionima sveta. Stoga je razvoj sorti pšenice koje su

11 prilagođene uslovima zaslanjenosti prioritet za globalnu sigurnost hrane. Procena

12 genotipova u realnim uslovima – in situ može pružiti odgovarajuće informacije o

13 performansama različitih genotipova u realnim uslovima i pomoći u identifikaciji

14 tolerantnih genotipova na salinitet. Da bi se obezbedila tačna procena dvadeset sedam

15 genotipova pšenice u različitim agroekološkim sredinama, zasnovan je ogled na dva

16 različita tipa zemljišta (solonjec i černozem) tokom dve vegetacione sezone. AMMI

17 analiza pokazuje da je faktor spoljašnje sredine imao najveće učešće (55,15%) u varijaciji

18 prinosa zrna, gde je dominantan uticaj imao tip zemljišta. Genotipovi Renesansa,

19 Harmonija i Bankut 1205 ostvarili su visok prinos zrna na oba tipa zemljišta. Međutim,

20 među navedenim genotipovima najveću toleranciju na salinitet ispoljio je genotip

21 Harmonija. Značajan deo interakcije genotipa i sredine (GEI; 25,89%) pokazuje da

22 postoji promena u rangiranju genotipova u različitim sredinama. Prema AMMI 1 biplotu,

23 genotipovi Renesansa i Harmonija su se odlikovali visokim prinosom zrna i visokom

24 stabilnošću. U agroekološkoj sredini Černozem 2015/2016, koja je imala najveći

1 doprinos interakciji, postignut je najveći prinos zrna, dok je u agroekološkoj sredini

2 Solonjec 2017/2018 ostvarena najveća stabilnost, ali i najniži prinos zrna. Prema AMMI 2

3 biplotu, genotip Harmonija je postigao visoku stabilnost u nepovoljnim uslovima sredine

4 Solonjec 2017/2018.

5 Ključne reči: aditivni efekti, multivarijacioni efekti, GEI, AMMI, stres saliniteta,

6 stabilnost

7 Tabela 1.
8
The share
Source of F– of total
Df SS MS p – value
variation value variation
(%)1
Total 323 913.2 2.83 - - 100
Treatmants 107 805.5 7.53 18.63** 0.000 88.21
Genotypes 26 102.0 3.92 9.71** 0.000 11.16
Environments 3 467.1 155.71 52.75** 0.000 51.15
Block 8 23.6 2.95 7.30** 0.000 2.58
Interactions 78 236.4 3.03 7.50** 0.000 25.89
IPCA1 28 146.9 5.24 12.98** 0.000 62.14
IPCA2 26 66.5 2.56 6.33** 0.000 28.13
IPCA3 24 23.0 0.96 2.37** 0.001 9.73
Residuals 0 0.0 - - - -
Error 208 84.1 0.40 - - -
9