Euphytica (2020) 216:184

https://doi.org/10.1007/s10681-020-02724-5 (0123456789().,-volV)( 0123456789().,-volV)

Genetic improvement of grain yield and bread-making quality of winter wheat over the past 90 years under the Pannonian Plain conditions

Milan Mirosavljevic´ . Vojislava Momcˇilovic´ . Dragan Zˇ ivancˇev . Vladimir Ac´in . Bojan Jockovic´ . Sanja Mikic´ . Verica Takacˇ . Srbislav Dencˇic´

Received: 18 March 2020 / Accepted: 27 October 2020 / Published online: 6 November 2020 Ó Springer Nature B.V. 2020

Abstract Studies on genetic gains in a historical set subunits N, 7 ? 9 and 5 ? 10 at the Glu-A1, Glu-B1 of wheat cultivars mostly refer to grain yield and the and Glu-D1, respectively, were the most prevalent in associated agronomical and physiological traits, while modern cultivars. Improvement of the main grain and less attention is paid to the improvement of wheat flour traits could be related with increased frequency quality. In order to analyze the breeding progress in of 7 ? 9 and 5 ? 10 subunits in modern wheat grain yield and with an emphasis on the main grain and cultivars. flour properties, twenty-five winter wheat cultivars released in the Southern Pannonian Plain between Keywords Breeding progress Á Flour attributes Á 1930 and 2015 were grown in three consecutive HMW-GS Á Protein seasons in (). The results from this study showed that grain yield increased linearly with an average rate of 45.5 kg ha-1 yr-1, implying that grain yield progress under conditions of the Southern Introduction Pannonian Plain have not reached the plateau. Protein and wet gluten content decreased significantly with Wheat (Triticum aestivum L.) is one of the most year of cultivar release, with a bi-linear trend. On the important staple grain crops in the world, produced on other hand, sedimentation value and gluten index 215 million hectares with an average grain yield (GY) increased in modern wheat cultivars, indicating of 3.43 t ha-1 (FAOSTAT 2020). Wheat breeding in improved protein quality. Also, the main flour traits the past century in many countries worldwide was related to stronger gluten and high sedimentation mainly directed at the GY improvement (Zhang et al. value, such as extensograph energy, farinograph 2016; Beche et al. 2018), while changes in grain resistance and alveograph deformation work, showed quality traits were a secondary breeding goal. How- improvement with year of cultivar release. Although ever, maintaining high grain quality with constant GY there were no significant changes in Gluten score, improvement is essential for human nutrition, as well as wheat milling and processing industry. Previous studies with a historical set of wheat M. Mirosavljevic´ (&) Á V. Momcˇilovic´ Á cultivars mainly dealt with the analysis of GY, main D. Zˇ ivancˇev Á V. Ac´in Á B. Jockovic´ Á agronomical and physiological traits providing infor- S. Mikic´ Á V. Takacˇ Á S. Dencˇic´ mation on the efficiency of selection (Sadras and Institute of Field and Vegetable Crops, Maksima Gorkog 30, Novi Sad 21000, Serbia Lawson 2011; Sanchez-Garcia et al. 2013; Kitonyo e-mail: [email protected] et al. 2017). Results from these studies reported that 123 184 Page 2 of 16 Euphytica (2020) 216:184

GY improvement predominantly related to the 20 years a significant number of cultivars from increase in grain number per unit area (Zhou et al. Western and Central Europe have been registered 2014; Valvo et al. 2018). Nevertheless, some recent and grown in Serbia and the surrounding Pannonian research showed an increase in grain weight in modern countries, being integrated into local breeding pro- cultivars compared to the older ones (Lopes et al. grams and affecting the genetic structure of the local 2012; Beche et al. 2014). Since wheat is mainly used in cultivars. Therefore, it is necessary to update and human diets, modern cultivars have to meet strict improve information about the current changes in quality requirements set by the food processing grain protein content and rheological properties of industry. Although some studies regarding breeding dough. The aim of this study was to analyze the progress in wheat analyzed certain quality traits, such breeding progress in GY and grain quality traits and to as protein content (PC) or hectoliter mass (Bilgin et al. assess the changes in the HMW-GS composition and 2016; Laidig et al. 2017), detailed investigations of the grain quality of winter wheat cultivars released in main grain and flour properties are scarce. Serbia and the surrounding countries from 1930 to Grain PC is an essential indicator of wheat end-use 2015. quality, and is under a significant effect of the environmental and genotypic variation (Hristov et al. 2010a). Protein quality is mainly determined by the Materials and methods composition of gluten molecules and its fractions, gliadin and glutenin. Variations in gliadin and glutenin Plant material composition affect gluten functionality and wheat flour baking properties (Delcour et al. 2012). Gener- A collection of 25 bread wheat cultivars released in or ally, dough extensibility is related to gliadin fractions introduced to Serbia between 1930 and 2015 was used (Branlard et al. 2001), while strength and elasticity of in this study. The selected cultivars represent main wheat dough is controlled by glutenin (Dhaka and historical and currently widely grown winter wheat Khatkar 2015; Zhang et al. 2018). Among glutenin cultivars in south Pannonian countries. Also, the subunits (high molecular weight glutenin subunits— studied cultivars have extensively been used in HMW-GS and low molecular weight glutenin sub- breeding activities and corresponded to the Serbian units—LMW-GS), HMW-GS play important roles in wheat breeding programs and programs from other the elasticity of dough (Shewry et al. 1992). The main breeding companies from Europe (, HMW-GS are encoded by Glu-A1 located on the long , , , and Hun- arm of the chromosome 1A, Glu-B1 on the chromo- gary). Year of release, country of origin and allelic some 1B, and Glu-D1 on the chromosome 1D (Payne differences in high molecular weight glutenin com- et al. 1981). Different compositions of HMW-GS have positions are reported in Table 1. a significant influence on dough strength and bread quality; therefore, information about historical Experimental field setup changes in HWG-GS is important for wheat breeders, since these are not commonly measured in the genetic Field trials were conducted over three growing seasons progress studies. (2016–2017, 2017–2018 and 2018–2019 marked as Studies on changes in wheat grain quality were 2017, 2018 and 2019, respectively) at the experimen- mainly limited to durum wheat (De Vita et al. 2007; tal field of the Institute of Field and Vegetable Crops, Subira et al. 2014) or to non-Pannonian climate zones, Novi Sad, Serbia (45°200N and 19°510E), which is a such as (Sanchez-Garcia et al. 2015), the United typical Pannonian environment. The soil at the site is a States of America (Fufa et al. 2005), or haplic chernozem aric (IUSS Working Group WRB (Guzma´n et al. 2017). Although wheat quality has 2014), and the main weather, soil physical and been an integral part of wheat breeding programs in chemical conditions of the 3 years are shown in the Pannonian plain, studies on genetic improvement Table 2. The experimental field had a long-term crop in quality traits in Serbia were conducted only on the rotation of cereals, maize and soybean. Sowing dates set of the cultivars released before 2000s (Dencic et al. were 10 October 2016, 13 October 2017 and 12 2007; Hristov et al. 2010b). However, in the previous October 2018, with target density of 550 plants per 123 Euphytica (2020) 216:184 Page 3 of 16 184

Table 1 The analyzed cultivars, country of origin, year of release and allelic composition at the high molecular weight glutenin subunits (HMW-GS) with Glu-score Cultivar Country of origin Year of release HMW-GS Glu-A1 Glu-B1 Glu-D1 Glu-score

Banatka Ukraine 1930 2* 7 ? 95? 10 9 Bankuti 1205 Hungary 1931 2* 7 ? 92? 12 7 San Pastore Italy 1940 1 20 2 ? 12 6 Mara Italy 1947 N 7 2 ? 12 5 Novosadska 1439 Serbia 1953 N 7 ? 95? 10 7 Bezostaya-1 Russia 1959 2* 7 ? 95? 10 9 Bacˇka Serbia 1964 2* 7 ? 95? 10 9 Libellula Italy 1965 1 20 2 ? 12 6 Sava Serbia 1970 1 7 ? 82? 12 9 Partizanka Serbia 1973 2* 7 ? 95? 10 9 Novosadska rana Serbia 1975 N 7 2 ? 12 5 Nizija Serbia 1979 2* 7 ? 92? 12 7 Jugoslavija Serbia 1980 2* 7 ? 95? 10 9 Lasata Serbia 1987 N 7 ? 85? 10 8 Skopljanka North Macedonia 1982 2* 7 ? 95? 10 9 Evropa 90 Serbia 1990 N 7 ? 92? 12 5 Pobeda Serbia 1990 2* 7 ? 95? 10 9 Renesansa Serbia 1994 N 7 ? 95? 10 7 Apache France 1998 2* 7 ? 93? 12 7 Simonida Serbia 2003 N 7 ? 92? 12 5 NS 40S Serbia 2006 N 7 ? 95? 10 7 Sirtaki France 2004 N 7 ? 82? 12 6 Ingenio France 2006 2* 7 ? 85? 10 10 NS Obala Serbia 2015 N 7 ? 95? 10 7 NS Mila Serbia 2014 N 7 5 ? 10 6 *is the name of HMW subunit and it is not related with the statistical significance square meter (recommended wheat sowing rates for Quantity and distribution of precipitation varied agroecological conditions of the south Pannonian significantly among the growing seasons, as is typical region). The experiment was arranged as a randomized for the Pannonian Plain. The total sum of precipitation complete block design with three replications. Plots from sowing to anthesis ranged from 223 mm in were 10 m long and 1 m wide, with 10 rows per plot. 2018/19 to 367 mm in 2017/18. During anthesis– In each growing season, soil samples were taken prior maturity period, total sum of precipitation was to plowing at the depths of 0–90 cm. Fertilizer 149 mm, 227 mm and 211 mm in 2016/17, 2017/18 combining nitrogen (N), phosphorus (P) and potas- and 2018/19, respectively. The mean daily tempera- sium (K) was applied prior to sowing to avoid N, P and tures from sowing to anthesis in 2016/17 were lower K deficit. In early February, additional nitrogen than in 2017/18 and 2018/19. On the other hand, mean fertilizer (ammonium nitrate—34% N) was top daily temperatures during grain filling period (anthesis dressed according to N-min analysis. As required in to maturity) were higher in 2016/17 and 2017/18 than spring, weeds, pests and diseases were controlled by in 2018/19. appropriate chemical application.

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Table 2 Basic description 2016/17 2017/18 2018/19 of the experimental site Coordinates 45°200N, 19°510E Altitude 84 m Soil classification Haplic chernozem aric soil Some properties of the soil

CaCO3 (%) 5.17 5.08 4.85

P2O5 (Egner–Riehm) (mg/100 g) 24.5 28.4 20.8

K2O (Egner–Riehm) (mg/100 g) 18.3 19.1 17.6 Humus (Turin) (%) 2.8 2.9 2.6 pH (KCl) 7.33 7.17 7.41 Fertilization -1 P2O5 (kg ha ) prior to sowing 60 55 70 -1 K2O (kg ha ) prior to sowing 60 55 70 N (kg ha-1) prior to sowing 60 55 70 N (kg ha-1) in spring 70 80 70 Temperature and precipitation during sowing-anthesis period Rainfall (mm) 280 367 223 Tmax (°C) 10.2 12.2 12.7 Tmin (°C) 0.9 3.2 3.4 Tmean (°C) 5.3 7.3 7.8 Temperature and precipitation during anthesis-harvest period Rainfall (mm) 149 227 211 *Tmax, Tmin and Tmean Tmax (°C) 26.3 27.1 23.7 are average maximum, Tmin (°C) 14.1 15.2 14.4 mean, and minimum daily Tmean (°C) 20.4 20.9 18.8 temperatures, respectively

Grain, flour and bread attributes half hour and milled by a Bu¨hler laboratory mill MLU- 202 (Uzwil, ). Falling number (FN) was All the plots were mechanically harvested at grain determined according to Hagberg with a Falling ripening (mid-June each growing season), and GY was Number 1800 instrument (Perten Instruments, Hud- expressed at 13% moisture level. From the harvested dinge, ) using the 107/1 method (ICC 2006). sample, thousand grain weight (g) was calculated as Sedimentation value (SV) of wheat samples were the mean weight of three sets of 200 grains from each determined according to AACC (2000) method replication. Grain nitrogen content was determined 56-62.01. Dough properties were tested by using a according to the Kjeldahl method by Kejltec 2300 farinograph (Brabender OHG, Duisburg, ) Analyzer Unit (Foss, Hilleroed, Denmark) by AACC according to the method 6369/653 (MSZ 1988). Also, (2000) method 46-10, and percent of protein content extensograph measurements (Brabender OHG, Duis- was calculated after multiplying Kjeldahl nitrogen by burg, Germany) were used according to the 114/1 5.7 and expressed on a grain dry weight basis. Wet method (ICC 2006). Flour strength (W, deformation gluten content (WG) and gluten index (GI) were energy) and the configuration ratio (P/L) were deter- determined by Glutomatic 2200 (Perten Instruments, mined with the alveograph (Chopin, Paris, France) Huddinge, Sweden) according to the ICC methods according to the 121 method (ICC 2006). 137/1 and 155. Before milling, the wheat grain samples were first tempered to 13.5% moisture content for 24 h. There- after, the wheat samples were tempered to 15.0% for

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HMW-GS allelic composition the agronomic significance and interpretability of the models tested. Samples with approximately 20 grains of each cultivar were analyzed for the determination of HMW-GS allelic composition. In order to define HMW-GS Results composition, total reduced seed proteins were sepa- rated by SDS-PAGE on 10% gel, in Tris–glycine HMW-GS allelic composition buffer. After the separation on the gel, the gels were stained by Commasie Brilliant Blue R-250 and fixed in The composition of HMW-GS of the winter wheat 7% acetic acid. The standard system of designating cultivars from this historical set varied according to glutenin loci, alleles and glutenin subunits was applied the breeding period and the country of origin. At the according to Payne and Lawrence (1983). The HMW- Glu-A1, there was a similar number of cultivars with N GS Glu-score was calculated according to Payne et al. and 2* subunits, while subunit 1 was registered in (1987), which indicated how genetic heritage affects three cultivars (San Pastore, Libellula and Sava). dough properties. The Glu-score for each cultivar, Nevertheless, since 1990 there has been an increase in which maximum value is 10, was calculated by frequency of N allele, since it was found in 70% of summing all individual HMW-GS scores. The allelic cultivars. At the locus Glu-B1, four alleles were found compositions at Glu-A1, Glu-B1 and Glu-D1 were that control 7 ? 9, 7 ? 8, 7 and 20 subunits. The obtained from published and breeders references subunit 7 ? 9 was predominant in the historical set at (Dencic et al. 2007; Hristov et al. 2010b; Lagrain the Glu-B1, since it was found in almost two-thirds of et al. 2013) and used to verify and complete the cultivars. Subunits 20, 7 and 7 ? 8 were found in two, obtained results. three and four cultivars, respectively. At the Glu-D1, subunit 5 ? 10 was most frequently observed in wheat Statistical analysis cultivars, while subunit 2 ? 12 was found in 10 cultivars and Apache was characterized by the com- The parameters of grain, flour and bread traits were bination 3 ? 12. There was no clear pattern of calculated for all three replicates and subjected to changes in Glu-score with year of cultivar release, analysis of variance (ANOVA) using Infostat (Di since cultivars with high and low score have been Rienzo et al. 2011). In order to partition the total identified among old, medium and modern cultivars. variation, ANOVA was performed using a generalized linear model (GLM) along with environments (grow- Component of variance ing seasons—GS), replications within environments, blocks within environments and their interactions. All Results from the Table 3 indicate that the effect of stated factors were treated as random effects. In order cultivar was significant for all traits. For most traits to evaluate the genetic progress for cultivars released (GY, PC, SV, WG, GI, SD, FR, R, EE, R/E and W), in different eras, values of all measured traits were cultivar had the greatest impact, accounting for [ analyzed as a function of the year of release with 50% of total variability in GY, PC, SV, GI, SD and SigmaPlot version 13.0 (Systat Software, San Jose, FR. Although the effect of the growing season was less CA). Linear or bilinear models, selected according to pronounced compared to the cultivar effect, all studied Akaike information criterion—(AIC) and the coeffi- traits were significantly affected by it. The greatest cient of determination (R2), were fitted in order to effect of growing season was observed in TGW establish the relationship between each studied traits (34.5%) and FN (45.5%). Also, the contribution of and the year of release. For bi-linear models the cultivar 9 growing season interaction was significant equation was: Y = A ? BX if X B C; and Y = A ? for all traits, with the highest influence on E and P/L. BC ? D(X- C) if X [ C; where X indicates the year of release, A is the intercept, B is the first slope Grain yield and weight representing the genetic progress, C indicates the year where a breaking point occurred and D indicates the Results from Fig. 1 showed that GY had a significant second slope of genetic progress. We also considered linear trend with year of cultivar release (R2 = 0.916). 123 184 Page 6 of 16 Euphytica (2020) 216:184

Table 3 Relative contributions to total sum of squares and significance level of source of variance for the studied traits of winter wheat cultivars released from 1930 to 2015 Traits Mean square % of SS GS G GS 9 GGSGGS9 G Error

GY (kg ha-1) 77,884,549** 12,930,541** 1,007,857** 28.2 56.2 8.8 6.8 TGW (g) 700.0** 43.3** 14.0** 34.5 25.6 28.5 11.4 PC (% d.m.) 3.54** 6.19** 7.89** 3.6 55.1 17.5 23.8 WG (%) 1271** 151** 43** 28.8 41 23.3 6.9 GI (%) 711** 1444** 578** 2.1 51 40.8 6.1 SV (ml) 1335** 215** 39** 25.9 50 18.3 5.8 FN (s) 498,151** 18,959** 13,943** 45.5 20,8 30.5 3.2 SD (BU) 1748** 4105** 1018** 2.2 62.1 30.8 4.8 FR (min) 543** 1052** 281** 1.5 62 34.6 1.8 E (cm2) 30,124** 3593** 2000** 24.4 34.9 38.9 1.8 R (BU) 468,738** 48,602** 18,161** 31 38.5 28.8 1.7 EE (mm) 28,662** 2580** 477** 36.9 39.9 14.8 8.4 R/EE 23.82** 3.07** 1.00** 27.8 43 28 1.2 W (10-4 J) 40,635** 11,607** 3823** 14.4 49.3 32.5 3.8 P/L 1.45** 0.38** 0.20** 13.3 42.1 43.9 0.7 GS growing season, G genotype, GS 9 G genotype by growing season interaction, GY grain yield, TGW thousand grain weight, PC protein content, WG wet gluten content, GI gluten index, SV sedimentation value, FN falling number, SD softening degree, FR quality group, E energy, R resistance, EE extensograph extensibility, R/E resistance/extensibility, W work of deformance, P/L tenacity/ extensibility ratio **Significant at the 0.01 probability level

The genetic progress in terms of GY was while after the inflection point trend in PC decline was 45.5 kg ha-1 yr-1. Average GY of wheat cultivars slower showing rate of 0.015% yr-1. Variation in PC varied from 7703 kg ha-1 in 2018 to 9627 kg ha-1 in was mostly related to difference between cultivars and 2017. GY of wheat cultivars varied from varied from 12.4% to 16.27%, with average PC of 5647 kg ha-1 to 10,599 kg ha-1. Thousand grain 13.8%. Averaged across cultivars, PC was similar in weight (TGW) showed a bi-linear response with year 2017 and 2019, while in 2018 cultivars had lower PC. of cultivar release. The genetic progress in TGW was The historical trend in wet gluten content (WG) also 0.1 g yr-1 until an inflection point in 1978. After showed a bi-linear trend. Similarly to PC, WG 1978, genetic progress in TGW reached plateau decreased at higher rate (0.243% yr-1) in cultivars showing an almost negligible decrease of released before 1964, and thereafter rate of WG 0.04 g yr-1. The average TGW of tested wheat reduction was lower (0.084% yr-1). The values of WG cultivars was 45.1 g, 45.3 g and 39.9 g in 2017, varied significantly between growing seasons, show- 2018 and 2019, respectively. Across growing seasons, ing highest WG in 2017 and lowest WG in 2018. WG TGW ranged from 39.5 g to 46.8 g. of the studied wheat cultivars ranged from 31.4% to 46.2%, with an average of 36.0%. Changes in gluten Grain quality traits index (GI) showed positive bi-linear trend of increase. Until 1975 cultivars showed increase in GI with rate of Grain protein content (PC) showed a bi-linear trend 0.463 yr-1, while GI improvement during the second with year of cultivars release, but in the opposite period (1975–2015) was 0.285 yr-1. Comparing direction to that observed in GY and TGW (Fig. 2). growing seasons, wheat cultivars achieved the highest Until 1982, PC was reduced at the rate of 0.04% yr-1, GI in 2019, and the lowest in 2018. The average GI

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Fig. 1 Environmental mean yield (a) and thousand grain significantly different at P = 0.05 by the Tukey test. Grain weight (b) of winter wheat cultivars grown in three growing yield (c) and grain weight (d) as functions of year of release seasons, means followed by the same letter were not across the growing seasons was 71.5, and among cultivars increased from 48.4 was significant influence of growing season on the FN, (Mara) to 91.3 (NS Obala). and the lowest FN (250 s) was recorded in 2017 and Sedimentation value (SV) showed a significant tri- the highest (413 s) in 2019. The FN varied between linear trend with year of cultivar release. The first 428 and 248 among cultivars. slope with genetic progress of 0.3667 ml yr-1 was observed until 1959 (Fig. 3). After this inflection Flour quality traits point, changes in SV decline to 0.0935 ml yr-1 until 1986. The genetic progress (0.03313 ml yr-1) during Degree of softening (SD) and farinograph resistance the last period 1999–2015 was also insignificant. (FR) have been analyzed in order to show the changes Between growing seasons, average SV was the highest in farinograph dough traits. According to Fig. 4,SD in 2019. The lowest SV was recorded in Banatka showed significant reduction from 1930 to 1970, but (22.35 ml), while the highest SW was reported in from 1970 onward it reached the plateau showing Ingenio (43.21 ml). The values of falling number (FN) insignificant decrease. Across cultivars, 2017 was varied significantly among the studied wheat geno- characterized by the highest average SD. The lowest types, ranged from 248 s to 428 s. There was a SD was recorded in Ingenio (37 BU) and the highest significant bi-linear relationship between FN and year SD was reported in Banatka (142 BU). Changes in FR of cultivars release. Values of FN decreased with rate had linear relationship with year of cultivar release. of 1.49 s yr-1 until 1994, and after that period FN The values of FR increased with rate of 0.044 yr-1. reached plateau showing insignificant decrease. There Among cultivars, FR ranged from 2.3 to 10.3 min,

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Fig. 2 Environmental mean protein content (a), wet gluten were not significantly different at P = 0.05 by the Tukey test. content (b) and gluten index (c) of winter wheat cultivars grown Protein content (d) wet gluten content (e) and gluten index (f)as in three growing seasons, means followed by the same letter functions of year of release across the growing seasons with an average value of 6.5 min. The values of FR 1930 and the inflection point, E and R had an increase ranged from 6.1 to 6.5 min in response to different of 0.707 cm2 yr-1 and 3.024 BU yr-1, respectively. environmental conditions. After inflection, E and R showed lower genetic Among different extensograph dough traits, energy progress. Among cultivars, E varied between 49 cm2 (E), resistance (R), extensibility (EE) and ratio and 137 cm2, while R ranged from 171 BU to 431 BU. between resistance and extensograph extensibility Although there was significant genotypic variation in (R/EE) were analyzed. Changes in E and R showed EE, results from regression analysis showed that there bi-linear trend with inflection point in 1972 and 1959, was no clear breeding effect on the EE. Also, EE respectively (Fig. 5). During the first period between varied between growing seasons, having highest

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Fig. 3 Environmental mean sedimentation value (a) and falling significantly different at P = 0.05 by the Tukey test. Sedimen- number (b) of winter wheat cultivars grown in three growing tation value (c) and falling number (d) as functions of year of seasons, means followed by the same letter were not release across the growing seasons average EE in 2018, and the lowest EE in 2017. (1.175) and the lowest in Banatka (0.379). These two Among cultivars, EE ranged from 127 to 209 mm, alveograph traits varied between growing seasons, and with an average value of 173.9 min. There was a the highest W and P/L were reported in 2019. positive linear relationship between R/EE and year of cultivar release with increase of 0.013 yr-1. Relationship between traits The deformation work (W) and alveograph ratio between tenacity and extensibility (P/L) are main Associations between the studied grain and flour traits alveograph traits according to the requirements of were explored with PCA analysis (Fig. 7). GY was trading sector (Fig. 6). The values of W had clear positively related with most alveograph, extensograph linear relationship with year of release, showing and farinograph traits (except EE). P/L, R/EE, TGW improvement trends of 0.803 10-4 Jyr-1. Among and R showed highest association with GY. Also, there cultivars, W varied from 110 to 251 10-4 J. Moreover, was close relationship between GI, SV, FR, E and W, P/L had tri-linear relationship with year of cultivar showing strong negative relationship with SD. On the release. According to the first slope of the relationship other side of the graph, PC and WG were closely P/L increased until 1950 with rate of 0.016 yr-1. After related with FN and EE. Most of the oldest cultivars the first inflection, values of P/L did not show genetic (released before 1970s) were placed on the left side of improvement until 1994 (second inflection), and after the graph, close to the vectors of PC and WG. Modern that P/L again started to rise significantly in modern cultivars separated on the graph in two groups. The cultivars. The highest P/L was recorded in Bacˇka first group with higher yielding cultivars (NS Mila, NS

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40 s and Sirtaki) was positioned close to the P/L, R/E Fig. 5 Environmental mean extensograph energy (a), resis- c and TGW, while the second group of modern cultivars tance (b), extensibility (c), and ratio between resistance and extensograph extensibility (d) of winter wheat cultivars grown (Ingenio and NS Obala) was characterized by higher in three growing seasons, means followed by the same letter values of GI, SV, E, W and FR. were not significantly different at P = 0.05 by the Tukey test. Extensograph energy (e), resistance (f), extensibility (g), and ratio between resistance and extensograph extensibility (H) as functions of year of release across the growing seasons Discussion

After the Second World War in former Yugoslavia, there was an essential need to increase wheat GY and middle of 2000s. Also, there were significant changes production in order to achieve self-sufficiency in in thousand grain weight (TGW), showing bi-linear domestic production and decrease wheat import. relationship with year of cultivar release. TGW -1 Increase of GY potential was confirmed with the increased with rate of 0.1 gr yr until 1978, after results from our study since breeding activities that progress in TGW reach the plateau. Therefore, resulted in significant GY improvement with rate of GY improvement during the first 50 years was more 45.5 kg ha-1 yr-1, indicating that rate of genetic gain related with TGW increase, and after 70 s with in winter wheat has not reached the plateau in southern increase in grain number per unit area. Pannonian Plain. Results from our study are similar to The main grain characteristics such as grain protein those reported by Mladenov et al. (2011) for south (PC) and wet gluten content (WG) showed bi-linear Pannonian plain winter wheat cultivars released up to trend of decline with year of cultivars release and

Fig. 4 Environmental mean degree of softening (a) and farino- significantly different at P = 0.05 by the Tukey test. Degree of graph resistance (b) of winter wheat cultivars grown in three softening (c) and farinograph resistance (d) as functions of year growing seasons, means followed by the same letter were not of release across the growing seasons 123 Euphytica (2020) 216:184 Page 11 of 16 184

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Fig. 6 Environmental mean degree of deformation work were not significantly different at P = 0.05 by the Tukey test. (a) and alveograph ratio (b) of winter wheat cultivars grown Deformation work (c) and alveograph ratio (d) as functions of in three growing seasons, means followed by the same letter year of release across the growing seasons negatively related with GY. The rate of PC (- 0.04% cultivars from Italy (San Pastore, Libellula and Mara) yr-1) and WG (- 0.243% yr-1) reduction was higher with a lower gluten index and their implementation in during the first phase, while during the second period local breeding programs. After the mid-1970s wheat after the inflection point reduction was smaller. GY and production in the former Yugoslavia became Reduction in PC and WG content is related to an higher and stable, and more attention in breeding increase in GY and TGW due to dilution of nitrogen programs was paid to the wheat quality (Dencic et al. compounds when carbohydrate synthesis is increased 2007), resulting in slower rate of PC and WG during photosynthesis (Jenner 1991). In contradiction reduction. On the contrary, values of gluten index to Dencic et al. (2007) and (Hristov et al. 2010a, b), our (GI) showed positive bi-linear trend of improvement results reported significant reduction in PC, while WG with year of cultivar release, indicating that gluten of decrease had with a higher rate from 1930 until the modern cultivars is stronger than in their older counter inflection point and lower rate after the inflection ones, as previously reported by Rakszegi et al. (2008) point. Also, rate of wheat PC decline was also reported and Hristov et al. (2010b). in other European countries (Peltonen-Sainio and The changes in sedimentation values (SV) and Peltonen 1994; Sanchez-Garcia et al. 2015; Bilgin falling number (FN) were significant in the previous et al. 2016; Laidig et al. 2017). From a historical period. In general, SV could be considered a good perspective, reported decrease in PC and WG during predictor of dough strength and bread volume (Pen˜a the period from 1930 until the first inflection point et al. 1990). SV increased from 1930 until 1959, and could be related to introduction of high yielding after inflection SV proceeded to rise with a lower rate,

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12 EE

8 FN W FR Novosadska rana E Evropa 90 Apache 4 Jugoslavija SV WG PC GI

Novosadska 1439 Ingenio

Bankut 1205 Mara Pobeda F2 (12.77 %) F2 (12.77 San Pastore Nizija Sava NS Obala 0 NS Mila Banatka Bezostaja 1 R Bačka Libbelula Lasta GY Skopjanka Sirtaki Simonida Partizanka NS 40S P/L

-4 SD Renesansa TGW R/EE

-8 -12 -8 -4 0 4 8 12 F1 (56.31 %)

Fig. 7 Biplot for measured grain and flour traits for 25 wheat cultivars (mean of 2017, 2018 and 2019) indicating that breeding activities during previous 1970, and after inflection point there were no signif- period maintained or even slightly improved protein icant changes in both farinograph traits. Due to a quality. The SV improvement was also reported in strong negative relationship between SD and GI (Kaur (Matus et al. 2012; del Pozo et al. 2014), et al. 2013), the improvement of protein quality Hungary (Rakszegi et al. 2008) and Germany (Laidig indirectly resulted in a decline in SD value. Among et al. 2017). Furthermore, FN decreased until 1994 different extensograph dough traits, energy (E), resis- with the rate of - 1.49 s yr-1. On the other hand, tance (R), extensibility (EE) and ratio between resis- Laidig et al. (2017) reported FN increase in German tance and extensograph extensibility (R/EE) were wheat (0.46%). Although FN was under a strong analyzed in this study. Changes in E and R showed bi- environmental control, in the modern wheat cultivars linear trend with inflection point in 1972 and 1959, values of this parameter are close to the limited values respectively. During the first period between 1930 and of the industry requirements, and further decrease the inflection point, E and R had increase of 0.707 cm2 (bellow 220–250) could lead to fewer financial yr-1 and 3.024 BU yr-1, respectively. After inflection benefits from milling sector. changes in E and R showed lower genetic progress of The impact of the breeding programs on the 0.206 cm2 yr-1, and 1.267 BU yr-1, respectively. rheological properties has particularly been high- Also, increase in extensograph E and R could be lighted in farinograph flour traits. Degree of softening related with stronger gluten and flour of modern (SD) had bi-linear relationship with year of cultivar cultivars, since more energy is required to stretch the release, while farinograph resistance (FR) was linearly flour (Hadnad¯ev et al. 2011). related. The values of SD decreased from 1930 until

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Furthermore, there was a significant effect of subunit with positive effect and 2 ? 12 subunit with breeding activities on alveograph dough properties negative effect on bread making quality (Popineau of wheat. During the past 90 years, the value of W et al. 1994) were present in 56% and 40% of cultivars, showed a linear trend of increase with year of release, respectively. Finally, the presence of 2 ? 12 subunit showing improvement trends of 0.803 10-4 Jyr-1. in Serbian cultivars, originating mostly from the Also, significant linear increase in W was reported in Italian cultivars, Bankut 1201 and 1205, was not Spain in the previous century, with higher rate of 2.74 recognized as a limiting factor for bread making 10-4 Jyr-1. Higher W increase in Spain could be quality, since cultivars with 2 ? 12 subunit could related to the presence of high number of Italian have high flour quality (Bedo¨ et al. 1995). cultivars in this country, characterized with increased values of W (Sanchez-Garcia et al. 2015). On the other hand, P/L had a tri-linear relationship with year of Conclusions cultivar release. From 1930 until 1950, P/L increased with rate of 0.016 yr-1, reaching a plateau that lasted In previous 90 years different trends have been until 1994. From 1994, increase in P/L values mainly observed for studied quality traits. Breeding progress results from the introduction of new cultivars with in grain yield in the southern Pannonian plain was higher values of tenacity (P). The values of W, FR and associated with a significant decrease in protein and E were highly related with SV and GI, and show no wet gluten content. Although there have been no clear relationship with PC and WG. Therefore, the changes in Glu-score of the studied historical set of increase of these main flour traits could be explained wheat cultivars in the previous 90 years, an increase in with improved gluten quality and predominance of GS sedimentation values, gluten index and most important 5 ? 10 at Glu-D1 and GS 7 ? 8 and 7 ? 9atGlu-B1 dough traits indicates that protein quality has in the past 20 years among the studied cultivars. improved. Improved gluten index and flour quality There were no clear changes in the Glu-score with could be related with increased frequency of 7 ? 9 year of cultivar release. Commonly, during wheat and 5 ? 10 subunits in modern cultivars released after breeding more attention is paid to the final bread 1990. quality than to the allelic composition, which is usually determined after the bread making quality Acknowledgements This study was carried out within the analysis. Considering Glu-A1 locus, cultivars with N project ‘‘Redesigning the exploitation of small grains genetic resources towards increased sustainability of grain-value chain or 2* subunit were present among old and modern and improved farmers’ livelihoods in Serbia and Bulgaria— ones. However, there was an increase in frequency of GRAINEFIT’’, PR-166-Serbia, funded by The Benefit-sharing N allele, related with lower sedimentation volume Fund of the International Treaty on Plant Genetic Resources for (Payne et al. 1987) among modern cultivars, since Food and Agriculture and COST Action ‘‘SOURDOugh biotechnology network towards novel, healthier and 1990 the presence of this subunit has increased to 70%. sustainable food and bioprocesses (SOURDOMICS)’’, This is significant increase in frequency of N allele CA18101. compared to the results of Dencic et al. (2007), who reported that 2* was the most frequent subunit in Authors’ contribution MM: methodology, writing—original draft. VM: conceptualization, formal analysis. DZˇ : cultivars released from 1993 until 2005. Also, culti- methodology, visualization. VA: investigation, methodology. vars with N allele are widely present in wheat cultivars BJ: writing—original draft, resources. SM: formal analysis. VT: in other European countries (Gregova´ et al. 1997; investigation. SD: validation, writing—review. Horvat et al. 2009; Filip 2018). Among the studied loci, Glu-B1 showed the highest variability and most Compliance with ethical standards studied wheat cultivars had 7 ? 9 and 7 ? 8 subunits Conflict of interest The authors declare that they have no characterized with positive effect on flour character- conflict of interest.. istic, as previously reported in different studies (Branlard et al. 2003; Gregova´ et al. 2004). Addition- ally, Hristov et al. (2010b) reported that GS subunit 7 ? 9 had the highest presence among modern wheat cultivars in Serbia. At the Glu-D1 locus, 5 ? 10 123 Euphytica (2020) 216:184 Page 15 of 16 184

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