© 2021 Al-Farabi Kazakh National University ISSN 1563-034Х; еISSN 2617-7358 Экология сериясы. №3 (68). 2021 https://bulletin-ecology.kaznu.kz
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IRSTI 34.35.15 https://doi.org/10.26577/EJE.2021.v68.i3.05 Y.A.Genievskaya1, 2 , S.S. Almerekova1, 2 , A.I. Abugalieva3 ,
V.A. Chudinov4 , S.I. Abugalieva1, 2 *
1Institute of plant biology and biotechnology, Kazakhstan, Almaty
2Al-Farabi Kazakh National University, Kazakhstan, Almaty
3Kazakh Research Institute of Agriculture and Plant Industry, Kazakhstan, Almalybak
4Karabalyk Agricultural Experimental Station, Kazakhstan, Karabalyk
*e-mail: [email protected]
GENOTYPE × ENVIRONMENT INTERACTIONS
IN GRAIN QUALITY TRAITS AND YIELD OF BARLEY GROWN IN KOSTANAY AND ALMATY REGIONS
Barley is one of the most important cereal crops in the world and the second cereal in Kazakhstan.
Traditionally barley is cultivated in the northern part of Kazakhstan, however, due to high genetic plastic- ity, barley has a great potential for adaptation in different climatic conditions. High levels of grain quality and yield are major indicators of good adaptation. In this study 357 spring barley cultivars and breeding lines from Kazakhstan and the USA were grown in two regions of Kazakhstan (Kostanay and Almaty regions). The collection was analyzed using seven grain quality and two yield-related traits, including grain protein, starch, lipids, ash, cellulose and moisture contents, plumpness, as well as thousand kernel weight and yield per square meter. A significantly large environmental impact was observed for thousand kernel weight and grain protein content. Grain starch, lipids content, and yield per square meter showed stability in two regions and were mostly controlled by the genotype. Correlations of high yield with high grain protein and low starch content were observed in both north and south-east regions. In general, the field trials assessment showed that barley cultivation in Kostanay region suits well malting-oriented breeding and in Almaty region for animal feed-oriented breeding. Based on the obtained results, the list of barley cultivars and promising breeding lines with high yield and grain quality was selected as the optimal breeding source in Kostanay and Almaty regions.
Key words: Hordeum vulgare L., malting barley, livestock feed barley, genotype × environment.
Ю.А.Гениевская1, 2, Ш.С. Альмерекова1, 2, А.И. Абугалиева3, В.А. Чудинов4, С.И. Абугалиева1, 2 *
1Өсімдіктер биологиясы және биотехнологиясы институты, Қазақстан, Алматы қ.
2Әл-Фараби атындағы Қазақ ұлттық университеті, Қазақстан, Алматы қ.
3Қазақ егіншілік және өсімдік шаруашылығы ғылыми-зерттеу институты, Қазақстан, Алмалыбақ а.
4Қарабалық ауыл шаруашылығы тәжірибе станциясы, Қазақстан, Қарабалық а.
*e-mail: [email protected]
Қостанай және Алматы облыстарында өсірілген арпаның дән сапасына және өнімділік белгілеріне генотип × қоршаған орта байланысының әсері
Арпа – әлемдегі ең маңызды дәнді дақылдардың бірі, ал Қазақстанда екінші маңызды дәнді дақыл болып табылады. Дәстүрлі түрде арпа Қазақстанның солтүстік бөлігінде өсіріледі, алайда оның жоғары генетикалық өзгергіштігінің арқасында арпаның әртүрлі климаттық жағдайларға бейімделу мүмкіндігі жоғары. Дәннің жоғары сапасы мен өнімділігі – жақсы бейімделудің негізгі көрсеткіштері болып табылады. Бұл жұмыста Қазақстанның екі облысында (Қостанай және Алматы облыстары) Қазақстан мен АҚШ-тан шыққан жаздық арпаның 357 сорты мен линиясы өсірілді. Коллекция дән сапасының жеті белгісі және өнімділіктің екі белгісі бойынша зерттелді, оның ішінде ақуыз, крахмал, май, күл, клетчатка мен ылғалдылық құрамы, дән натурасы, сонымен қатар 1000 дәннің салмағы мен шаршы метрге шаққандағы өнімділік сияқты екі өнімділік белгісі зерттелді. Қоршаған орта 1000 дәннің салмағы мен дәндегі ақуыз құрамына едәуір әсер етті. Крахмал, май мөлшері және шаршы метрдегі өнімділік екі аймақта тұрақтылықты көрсетті және генотипке тәуелділігі анықталды. Жоғары өнімділіктің жоғары ақуыз құрамымен және астық құрамындағы төмен крахмалмен корреляциясы солтүстікте де, оңтүстік-шығыста да байқалды. Жалпы, далалық сынақтарды бағалау Қостанай облысында өсірілген арпа сыра қайнату бағытында, ал Алматы облысында жемге деген қажеттілік үшін өсіруге қолайлы екенін көрсетті. Алынған нәтижелер бойынша Қостанай және Алматы облыстарында өсіруге донор
Y.A.Genievskaya et al.
ретінде жоғары өнімділігі мен астық сапасына ие арпаның перспективті сорттары мен линиялары таңдалды.
Түйін сөздер: Hordeum vulgare L., сыра қайнатуға арналған арпа, жемдік арпа, генотип × қоршаған орта.
Ю.А. Гениевская1, 2, Ш.С. Альмерекова1, 2, А.И. Абугалиева3, В.А. Чудинов4, С.И. Абугалиева1, 2 *
1Институт биологии и биотехнологии растений, Казахстан, г. Алматы
2Казахский национальный университет им. аль-Фараби, Казахстан, г. Алматы
3Казахский научно-исследовательский институт земледелия и растениеводства, Казахстан, пос. Алмалыбак
4Карабалыкская сельскохозяйственная опытная станция, Казахстан, пос. Карабалык
*e-mail: [email protected]
Влияние взаимодействия генотип × среда на признаки качества зерна и урожайности ячменя, выращенного в Костанайской и Алматинской областях Ячмень – одна из важнейших зерновых культур в мире и вторая по значимости среди зерновых в Казахстане. Традиционно ячмень выращивают в северной части Казахстана, однако, благодаря высокой генетической пластичности ячмень обладает большим потенциалом для адаптации в различных климатических условиях. Высокие качество зерна и урожайность являются основными показателями хорошей адаптации. В данной работе 357 сортов и линий ярового ячменя из Казахстана и США были выращены в двух регионах Казахстана (Костанайской и Алматинской областях). Коллекция были исследована по семи признакам качества зерна и по двум признакам урожайности, включая содержание в зерне белка, крахмала, жиров, золы, клетчатки и влаги, натуру зерна, а также массу 1000 зерен и урожайность на квадратный метр.
Наибольшее значительное влияние среда оказала на массу 1000 зерен и содержание белка в зерне. Уровни крахмала, жиров и урожайности на квадратный метр показали стабильность в двух регионах и в большей степени зависели от генотипа. Корреляции высокой урожайности с высоким содержанием белка и низким содержанием крахмала в зерне наблюдались и на севере, и на юго-востоке. В целом, оценка полевых испытаний показала, что ячмень, выращенный в Костанайской области, более пригоден для селекции в пивоваренном направлении, в то время как в Алматинской области – для кормовых нужд. Основываясь на полученных результатах, были отобраны перспективные сорта и линии ячменя с высокими урожайностью и качеством зерна в качестве доноров для селекции в Костанайской и Алматинской областях.
Ключевые слова: Hordeum vulgare L., пивоваренный ячмень, кормовой ячмень, генотип × среда.
Abbreviations
ANOVA – analysis of variance, GAC – grain ash content, GCC – grain cellulose content, GE – geno- type × environment interaction, GLC – grain lipid content, GMC – grain moisture content, GPC – grain protein content, GPL – grain plumpness, GSC – grain starch content, KAES – Karabalyk agricultural exper- imental station, KRIAPI – Kazakh research institute of agriculture and plant industry, TKW – thousand kernel weight, YM2 – yield per m2.
Introduction
Barley (Hordeum vulgare L.) is a cereal crop with an important role in the World food security and economics. Due to the high genetic plasticity and adaptability of this species, it is successfully cultivated in all regions of the World. In Kazakh- stan, barley takes 18.25 % of the total cereal produc- tion (during the season 2019 / 2020), which is the
second place in the country after wheat [1]. Along with the internal market, Kazakhstan’s barley is ex- ported abroad and makes Kazakhstan the 7th largest barley exporter in 2019 / 2020 [2].
Traditionally, in Kazakhstan, commercial culti- vation of spring barley is focused in Akmola and North Kazakhstan regions. Still, in the season 2019 / 2020, Kostanay and Almaty regions produced 12.0
% and 12.7 % of the total spring barley in Kazakh- stan, respectively [1], demonstrating the good po- tential to compete with the leading regions. Both regions have a list of barley cultivars specifically developed for their soil and climatic conditions. For example, according to the State Register of Breed- ing Achievements of Kazakhstan 2020 [3], there are 31 spring barley cultivars of local and foreign breed- ing officially approved for the Almaty region and 17 cultivars for the Kostanay region [3]. Also, some of these cultivars demonstrate high plasticity with good yield performance in several other regions. For instance, cultivar “Arna” was approved for Almaty,
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Genotype × environment interactions in grain quality traits and yield of barley grown in Kostanay and Almaty regions
East Kazakhstan, Zhambyl, Turkestan, Akmola, Kostanay, and North Kazakhstan regions [3]. It was developed by KRIAPI and used for many studies in different regions of Kazakhstan [4-6]. This ex- ample shows the possibility of introducing outside bred barley cultivars to their successful use in local breeding processes.
Usually, barley is cultivated for three major di- rections – malting and brewing, livestock feed, and for the state purchases and deliveries. In Kazakh- stan, barley is mainly used for livestock feed (more than 43 %) and about 37 % for the food industry and malting. Depending on the breeding direction, there is a difference in requirements for the quality of barley grain. For example, raw protein content in barley grain used for malting is recommended to be less than 12 % [7], while barley grain for the live- stock feed has to contain more than 13 % of raw pro- tein [8]. Due to the long history of livestock in Ka- zakhstan, most modern local barley cultivars were developed for the feed. However, there are three new cultivars – Arna, Asem, and KazSuffle-1 – de- veloped for malting with good grain quality traits.
Nevertheless, the ratio of local malting barley culti- vars in Kazakhstan to cultivars for livestock feed is extremely small.
It was shown that important traits of barley grain quality are under polygenic control, including protein [9], starch [10], and cellulose content [11].
Still, the impact of environment and GE interaction significantly influences grain quality traits, some- times even more than the genotype. Particularly, the effect of GE was observed for diastatic activity and malt extract of barley [12]. At the same time, studies reveal low or insignificant GE interaction for malting quality of barley, but a remarkable influ- ence on grain yield [13]. Individual barley cultivars and breeding lines were studied for grain yield and quality in Kostanay region [14], Central Kazakhstan [15], and Turkestan region [16]. Significant effect of the environment on barley grain yield and yield- related traits was also discovered in different regions of Kazakhstan [17-18], where the average yield in the Kostanay region was significantly higher than in the Almaty region. However, a comprehensive com- parative analysis of barley quality traits in different regions of Kazakhstan has not been previously con- ducted.
Usually, high barley grain yield is negatively correlated with good malting quality traits, which creates difficulties for breeders and agronomists.
The development of barley cultivars with the opti- mal combination of grain yield and quality for dif- ferent growing regions is an important modern prob-
lem. In this regard, the study of yield and quality traits in modern cultivars and future promising lines of barley, considering the climate in a particular re- gion of Kazakhstan, has a high priority.
Materials and methods
Barley accessions used in the study. The set of barley accessions included 267 two-rowed spring barley cultivars and breeding lines from the USA and 90 two-rowed spring barley cultivars and breed- ing lines from Kazakhstan. The USA’s part of the set was represented by accessions from Montana State University, Washington State University, Utah State University, Small Cereal Collection of the USDA held in Aberdeen, Idaho, and one private com- pany, Busch Agricultural Resources, a division of the Anheuser-Busch Corporation [17]. Accessions from Kazakhstan were obtained from 6 agricultural institutions: Aktobe Agricultural Experimental Sta- tion (Aktobe region), Karabalyk Agricultural Ex- perimental Station (Kostanay region), Karaganda Agricultural Experimental Station (Karaganda re- gion), Kazakh Research Institute of Agriculture and Plant Industry (Almaty region), Kazakh Research Institute of Rice-growing (Kyzylorda region), and Krasnovodopad Agricultural Experimental Station (Turkestan region) [17]. Each institution provided barley accessions adapted to different regions of Ka- zakhstan.
Field experiment. All 357 accessions were grown in two environments – in the fields of KAES (Kostanay region) and KRIAPI (Almaty region) during the 2020 season (Figure 1). Each individual accession was sowed in 1 m2 plot in two indepen- dent replications. In order to reveal relations be- tween grain quality and yield, TKW (g) and YM2 (g/m2) as yield-related traits were measured in both environments for each individual accession in two replications. Cultivars “Ubagan” and “Asem” were chosen as standard check cultivars for KAES and KRIAPI, respectively. “Ubagan” is one of the best livestock feed cultivars, while Asem was developed for its malting qualities. Both cultivars are regis- tered and recommended for cultivation in respective regions by the State Register of Breeding Achieve- ments of Kazakhstan [3].
Analysis of grain quality traits. Grains of all 357 accessions were studied using seven ma- jor quality traits in the Laboratory of grain quality (KRIAPI). The set of barley accessions was studied for the content of raw ash (GAC, %), raw cellulose (GCC, %), raw lipid (GLC, %), raw grain moisture (GMC, %), raw protein (GPC, %), raw starch con-
Y.A.Genievskaya et al.
tent (GSC, %), and grain plumpness (GPL, g/L).
GPL was measured according to the GOST [19].
Remaining quality traits were determined by NIRS
DS 2500 Grain Analyzer (FOSS, Hillerød, Den- mark), with a calibration supplied by the manu- facturer.
Figure 1 – Meteorological data by barley growth stages in KAES (Kostanay region) and KRIAPI (Almaty region)
Statistical analysis. ANOVA test was per- formed to determine the effects of genotype, envi- ronment, and genotype × environment on the stud- ied traits. Pearson correlation analysis was carried out to reveal interrelations among grain quality traits and their connection with the yield. Both analyses and traits bar plots were performed using R statisti- cal platform [20].
Results and discussion
Quality traits and yield in two regions. Seven barley grain quality traits and two yield-related traits were studied in two climatic conditions of Kazakh- stan – north (KRAES) and south-east (KRIAPI).
The obtained results showed moderate differences in GAC, GCC, GMC, GPC, and TKW, small differ- ences in GPL and YM2, and almost no difference in GLC and GSC between the two regions (Figure 2).
Barley accessions grown in KAES had demonstrat- ed larger values of GAC and GPL, while accessions from KRIAPI had higher levels of GCC, GMC, GPC, and higher TKW and YM2. A significant dif- ference in temperature between the two regions was observed only at the period between sowing and til- lering; the amount of precipitations was significant- ly higher in KRIAPI (Figure 1).
As mentioned earlier, barley production in Kazakhstan is oriented into three basic directions (livestock feed, malting, and state purchases/deliv- eries of the grain). Each direction has its specific requirements for grain quality. Thereby, studied
traits should be compared between regions and state standards (GOST) according to these directions.
According to the GOST R 53900-2010 [8], barley grain for the 1st class livestock feed has to demon- strate high raw protein content (> 13 %), but low raw cellulose (< 7 %) and ash (< 2 %) content. Bar- ley grown in KRIAPI fitted better to quality require- ments for livestock feed grain in combination with a relatively higher yield. It makes it possible to sug- gest that in 2020 south-east Kazakhstan had more favorable conditions for the cultivation of barley for livestock feed. The GOST 28672-90 [21] for the state purchases and deliveries of the 1st class barley grain requires < 19 % moisture content and grain plumpness > 630 g/L. As for the GOST 5060-86 [7], regulating grain quality of barley for malting re- quires < 12 % of raw protein and < 15 % of moisture content in the 1st class grain. The concentration of raw starch between 60 % and 70 % is not mentioned in the GOST 5060-86, but it is considered as an op- timum for malting.
A stable combination of high grain yield and ap- propriate grain quality traits is the primary goal for all breeders. However, often increasing yield leads to undesirable changes in grain quality of barley.
Correlation test among studied traits in two regions showed significant positive correlations between TKW and YM2, TKW and GPC, and between GMC and GSC; and negative correlations between GSC and GLC and between GSC and GPC observed in both regions (Figure 3). A positive correlation be- tween yield and protein content in barley grain
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Genotype × environment interactions in grain quality traits and yield of barley grown in Kostanay and Almaty regions
was also previously observed in Kyzylorda region [22], where the same barley accessions were stud- ied. Still, if the combination of high yield (TKW, YM2) and high GPC is good for livestock feed grain, a high protein level decreases the quality of the brewing end-product – beer [23]. Stable nega- tive correlation between GSC and GPC was in ac- cordance with previous studies [24-26], where the
authors conclude that barley GPC increases as a result of the suppression of starch synthesis dur- ing the kernel development. Low starch in barley grain results in a low concentration of simple sug- ars in the malt, which leads to poor fermentation [27]. Therefore, it is important to find the optimal combination of GPC, GSC levels, and the yield for malting cultivars.
Figure 2 – comparison grain quality and yield-related traits of barley grown in KAES (Karabalyk agricultural experimental station, Kostanay region) and KRIAPI
(Kazakh research institute of agriculture and plant industry, Almaty region)
As for the other traits, in this studied collec- tion, a positive correlation between GAC and GCC, and a negative between GPC and GCC observed in KRIAPI confirms the suggestion that south-east Kazakhstan had more favorable conditions for the cultivation of barley for livestock feed.
In general, considering the abovementioned requirements for the entire collection, barley grown in KAES in 2020 is more suitable for bar- ley production for malting, grain state purchases, and export. These observations confirmed the leading role of north Kazakhstan in barley pro- duction for the malting and food industry. At the same time, conditions of south-east Kazakhstan
suits better for the cultivation of barley for live- stock feed.
The impact of genotype and environment on barely grain quality and yield. For a better under- standing of genotype’s and environment’s role in barley grain quality traits and yield, ANOVA was performed. Based on ANOVA results, all studied traits can be divided into three groups: those where the genotype explained the maximum effect – GPL, GSC, and YM2; those where the environment dem- onstrated the largest effect – GPC, GCC, and TKW;
and those where the impacts of genotype, environ- ment, and G×E were less than the residuals – GMC, GLC, and GAC (Table 1).
Y.A.Genievskaya et al.
Figure 3 – Plots of Pearson correlation coefficients (r) among barley grain quality traits and yield in KRAES, Kostanay region (A) and KRIAPI, Almaty region (B). Cells with significance at
P < 0.05 are highlighted in color. Red color denotes negative correlation, blue color – positive. Color intensity increases with the decreasing of P
Table 1 – ANOVA results for grain quality and yield-related traits of barley grown in two regions of Kazakhstan
Trait Factor df SS MS Var. F p Var. G
(%) Var.
E (%) Var. G×E (%)
GPL G 358 620053 1732 442.26 3.06 < 2E-16
39.0 11.4
E 1 180845 180845 128.99 319.32 < 2E-16 24.9
G×E 351 395344 1126 281.99 1.99 1.10E-14
Res. 691 391341 566 279.13
GMC G 358 89.4 0.25 0.06 1.21 0.0181
23.2 31.7
E 1 122.03 122.03 0.09 591.05 < 2E-16 7.9
G×E 351 30.38 0.09 0.02 0.42 1.0000
Res. 691 142.67 0.21 0.10
GPC G 358 247.1 0.7 0.18 2.67 < 2E-16
21.6 50.6
E 1 578.7 578.7 0.41 2240.24 < 2E-16 12.2
G×E 351 139.6 0.4 0.10 1.54 9.59E-07
Res. 691 178.5 0.3 0.13
GLC G 358 78.98 0.221 0.06 1.42 0.0001
31.2 1.5
E 1 3.89 3.891 0.00 24.97 7.37E-07 24.7
G×E 351 62.61 0.178 0.04 1.15 0.0694
Res. 691 107.66 0.156 0.08
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Genotype × environment interactions in grain quality traits and yield of barley grown in Kostanay and Almaty regions
Trait Factor df SS MS Var. F p Var. G
(%) Var.
E (%) Var. G×E (%)
GCC G 358 116.45 0.33 0.08 1.52 1.76E-06
22.1 31.9
E 1 168.19 168.19 0.12 785.81 < 2E-16 17.7
G×E 351 93.28 0.27 0.07 1.24 0.0089
Res. 691 147.68 0.21 0.11
GAC G 358 36.84 0.1 0.03 1.63 2.62E-08
25.4 23.4
E 1 33.9 33.9 0.02 537.74 < 2E-16 21.1
G×E 351 30.56 0.09 0.02 1.38 0.0002
Res. 691 43.56 0.06 0.03
GSC G 358 889.5 2.48 0.63 2.72 < 2E-16
43.6 3.1
E 1 63.1 63.15 0.05 69.17 4.81E-16 22.3
G×E 351 455 1.3 0.32 1.42 0.0001
Res. 691 630.9 0.91 0.45
TKW G 358 19329 54 14.02 6.04 < 2E-16
13.6 76.7 5.3
E 1 108817 108817 78.91 12134.63 < 2E-16
G×E 351 7516 22 5.45 2.41 < 2E-16
Res. 691 6017 9 4.36
YM2 G 358 19907881 55764 14436.46 2.09 < 2E-16
36.2 12.5
E 1 6873994 6873994 4984.77 257.50 < 2E-16 18.7
G×E 351 10273534 29522 7449.99 1.11 0.1370
Res. 691 17912606 26695 12989.56
Notes: df – degree of freedom, SS – sum of squares, MS – mean square, Var. – variance, G – genotype, E – environment, G×E – genotype × environment, Res. – residuals.
Continuation of table 1
The largest environmental effects were observed for TKW (76.7%, P < 2E-16) and for GPC (50.6 %, P < 2E-16) (Table 1). Both traits are complex, under polygenic control [28, 9], and demonstrates posi- tive correlation (Figure 3). However, studies were revealing a strong impact of the environment on bar- ley kernel weight [29] and the protein content [30- 31]. The significant difference in soil and weather conditions of the two regions gave a large variance in TKW and GPC between north and south-east Kazakhstan. TKW and GPC are very important for both livestock feed and malting barley; therefore, both genetic and environmental factors should be considered by breeders. The levels of three other important barley traits – GPL, GSC, and YM2 – were mostly influenced by the genotype (P < 2E-16) with 39.0 %, 43.6 %, and 36.2 % of the variance, respectively (Table 1). These traits are also complex with genetic and environmental control [31-34]. The stability of these traits between regions is a good source for the development of multi-environmental barley cultivars for different regions of Kazakhstan.
Significant moderate impact of G×E was detected for GPL (24.9 %, P < 1.10E-13), GAC (21.1 %, P <
0.001), and for GSC (22.3 %, P < 0.01). However, for all of the studied traits, the effect of G×E was not the largest one (Table 1).
The selection of barley accessions with promising grain quality and yield. Each region of Kazakhstan has a list of barley cultivars rec- ommended for cultivation called the State Regis- ter of Breeding Achievements of Kazakhstan [3].
This list includes cultivars of local and foreign breeding, and it is updated every year. For ex- ample, barley cultivar “Ubagan” is recommended for Kostanay region. However, according to the results of this study, there were accessions with good quality of the grain demonstrating higher yield than the standard “Ubagan”. The list of promising accessions for livestock feed barley in the Kostanay region includes six cultivars and breeding lines from Kazakhstan and four acces- sions from the USA (Table 2). In the Almaty region, the list of promising accessions for live-