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<article xmlns:xlink="http://www.w3.org/1999/xlink">
  <front>
    <journal-meta />
    <article-meta>
      <title-group>
        <article-title>Evaluation of valuable agronomic traits of spring triticale in a long-term breeding trial</article-title>
      </title-group>
      <contrib-group>
        <contrib contrib-type="author">
          <string-name>Irina G. Grebennikova</string-name>
          <xref ref-type="aff" rid="aff0">0</xref>
        </contrib>
        <contrib contrib-type="author">
          <string-name>Anna F. Cheshkova</string-name>
          <xref ref-type="aff" rid="aff0">0</xref>
        </contrib>
        <aff id="aff0">
          <label>0</label>
          <institution>Siberian Federal Scientific Center of Agro-BioTechnologies of the Russian Academy of Sciences</institution>
          ,
          <addr-line>Krasnoobsk, Novosibirsk region</addr-line>
          ,
          <country country="RU">Russia</country>
        </aff>
      </contrib-group>
      <fpage>454</fpage>
      <lpage>462</lpage>
      <abstract>
        <p>The analysis of data of long-term breeding trials of spring hexaploid triticale, used for forage and grains, was carried out using mathematical methods and developed software. The valuable agronomic traits of the ideal variety have been determined. The possibility of purposeful breeding for adaptability to the conditions of the region when creating new high-yielding varieties is shown.</p>
      </abstract>
      <kwd-group>
        <kwd>eol&gt;Spring triticale</kwd>
        <kwd>plant breeding</kwd>
        <kwd>valuable agronomic traits</kwd>
        <kwd>long-term breeding data</kwd>
        <kwd>computer program</kwd>
      </kwd-group>
    </article-meta>
  </front>
  <body>
    <sec id="sec-1">
      <title>1. Introduction</title>
      <p>
        The main strategy in the breeding of cereals is to create varieties with high productivity potential,
which would be realized by at least 70–80% under actual production conditions. New varieties
have the greater value, the more optimally they combine the important biological, economic and
technological properties. The beneficial features of the variety can be seen only under certain
growing conditions that ensure the realization of its potential productivity. New varieties must
combine adaptability to certain agro-climatic zone with all other valuable agronomic traits.
When creating a new variety, it is necessary to take into account the agro-climatic zone for
which the variety is being created, soil cover, meteorological factors, the spread of diseases
and plant pests, and the features of agricultural technology. The breeder also needs to identify
critical weather and climatic factors afecting the cereals. All this determines the main direction
of crop breeding [
        <xref ref-type="bibr" rid="ref1 ref2">1, 2</xref>
        ].
      </p>
      <p>
        One of the hopeful crops for Western Siberia is triticale, highly appreciated in many countries
including Russia for its versatility and high biological potential. Triticale has a high adaptability
to growing conditions, a greater yield on depleted soils compared to wheat, better grain quality
than rye and higher protein content in grains. Due to rye genome triticale is immune to
many cereal diseases such as powdery mildew, common bunt, loose smut. Its seeds do not
require fungicide treatment before sowing which avoids high production costs and preserves
the ecological balance of soils. Pests damage triticale less than wheat, due to the presence of
biological defense mechanisms [
        <xref ref-type="bibr" rid="ref3">3</xref>
        ].
      </p>
      <p>
        A characteristic feature of triticale is that the earing, flowering and ripening stages occur
later and last longer than in wheat. Late earing of triticale is an advantage for forage production
compared to rye. The green biomass yield of forage triticale is 2 times higher than that of wheat.
Forage triticale surpasses winter rye in protein, carotenoid and sugar content. Lignification in
triticale is slower than in rye, which preserves the feeding quality of the stems after flowering.
As a result, green biomass is better eaten by animals and provides higher weight gain. The use
of triticale both for fodder and grains in some cases has an advantage over the use of wheat,
corn, barley and sorghum. Thanks to these advantages, triticale significantly reduces the cost
and diversifies the production of high-quality feed and food grains, and enables rational use of
the available soil and climatic resources [
        <xref ref-type="bibr" rid="ref4 ref5">4, 5</xref>
        ]. All these qualities define triticale as a crop of
low environmental risk.
      </p>
      <p>
        Despite a number of beneficial features, spring triticale in Western Siberia is a new and poorly
studied crop. Agro-technical methods of triticale cultivation were developed mainly for the
European regions of Russia or for foreign countries and therefore are not always acceptable for
Western Siberia. Practical experience and scientific research show that foreign varieties have
weak local adaptation [
        <xref ref-type="bibr" rid="ref6">6</xref>
        ]. An increase in the sowing area of triticale and widespread production
of this crop is possible only by creating varieties highly adapted to local agroecological conditions,
combining high productivity and high field resistance to various phytopathogens.
      </p>
      <p>The objective of this study is to evaluate morphological and agronomic traits and ecological
adaptability of spring triticale used for forage and grain in a long-term breeding trial.</p>
    </sec>
    <sec id="sec-2">
      <title>2. Materials and methods</title>
      <p>Desirable agronomic traits can be identified by analyzing varieties with high adaptive potential,
which manifests itself in unfavorable environments, grown for a long time in a specific
agroclimatic zone. Adaptation of genotypes to local conditions occurs due to the heterozygosity of
triticale and due to the bufering efect of the genetic variety. The advantage of evolutionary
selection is that at the end of the trial we have a population enriched with valuable and promising
genotypes, which can become material for individual selection. This method helps to determine
the desired biotype or model of the ideal variety, towards which the eforts of breeders should
be directed.</p>
      <p>A field trial was conducted in Novosibirsk region, Russia in 2018–2020 at the fields of SFSCA
RAS and SRIPCB (branch of ICG SB RAS). Experiment included three varieties of common
wheat and four breeding lines of spring hexaploid triticale (Table 1).</p>
      <p>
        The triticale lines studied in this work are intended for use grains in food and animal feed.
When evaluating valuable agronomic traits of spring triticale, comparison was made with wheat
varieties of Siberian selection of diferent ripeness groups. Since at the time of the field trials
there were no data on the triticale variety Timur entered in the register of breeding achievements
in the West Siberian region in 2020 [
        <xref ref-type="bibr" rid="ref7">7</xref>
        ]. Sowing was carried out manually in open ground in
four replications, 80 grains in a row with a feeding area of 0.3 m2 .
      </p>
      <p>The following traits were registered: plant height, spike length, number of spikelets per
spike, ear density, grains weight of an ear, number of grains per spike, weight of 1000 grains,
dynamics of plant growth and phenological phases duration. Biological control of plant revealed
diferences in growth, development and grains formation of spring wheat and triticale, which
are influenced by environmental conditions (day length, the spectrum of solar insolation,
temperature, air humidity and soil moisture).</p>
      <p>Yield is the total manifestation of the productivity and adaptability potentials of the studied
genotypes. The potential yield was determined from the analysis of the productivity of the five
best plants selected from each replication. The actual yield was determined based on the yield
of the plot.</p>
      <p>To assess the environmental adaptability of varieties, the following agrotechnical methods
were used: 1) field trials in diferent soil conditions; 2) field trials in three diferent length of
daylight.</p>
      <sec id="sec-2-1">
        <title>Resistant to lodging, moder</title>
        <p>ately drought-resistant.
Moderately susceptible to common
bunt. Strongly susceptible to
brown and stem rusts, powdery
mildew.</p>
        <p>Resistant to lodging,
moderately drought-resistant.
Moderately susceptible to brown rust
and septoria. Strongly
susceptible to loose smut.</p>
        <p>Resistant to lodging, drought
tolerant. Moderately
susceptible to septoria. Strongly
susceptible to loose smut, brown rust
and powdery mildew.</p>
      </sec>
      <sec id="sec-2-2">
        <title>Medium-dense ear, with long</title>
        <p>awns. Resistant to lodging.
Susceptible to ergot fungi.</p>
        <p>Plants with medium awn length
were selected. Dense ear. High
quality grains. Resistant to
lodging. Drought tolerant. Powdery
mildew resistant.</p>
        <p>Dense ear with long awns. High
quality grains. Resistant to
lodging. Drought tolerant. Powdery
mildew resistant.</p>
        <p>Obtained from diallel crosses.</p>
        <p>Low stem, Resistant to lodging.</p>
        <p>Dense ear, awnless. Powdery
mildew resistant.</p>
        <p>During the period of field trials there was quite a large variety of typical for Western Siberia
weather conditions, particularly in seasonal distribution and sum of air temperature and
precipitation from May to August. This allowed a thorough study the genotypes.</p>
        <p>
          The analysis of long-term breeding data on early maturity, stress resistance and productivity
was carried out using the following mathematical methods and software tools. Multivariate
correlation analysis was used to assess the relationship between various traits of the studied
samples [
          <xref ref-type="bibr" rid="ref8">8</xref>
          ]. The Ward’s method of cluster analysis was used to compare the studied samples by
a complex of traits and to select a group of plants that most closely matched the model of ideal
variety [
          <xref ref-type="bibr" rid="ref9">9</xref>
          ]. The best samples were selected by evaluation of the plant breeding index, which was
defined as the weighted sum of normalized productivity traits according to the formula [
          <xref ref-type="bibr" rid="ref10">10</xref>
          ]:
where  is the number of traits;  — expert weighting coeficients satisfying the condition

 = ∑︁  · ,
        </p>
        <p>=1

∑︁  = 1.</p>
        <p>=1
 =</p>
        <p>− min()
max() − min()
,
(1)
(2)
Here  ( = 1, . . . , ) are normalized values of traits:
where max(), min() — the maximum and minimum values of the -th trait.</p>
        <p>
          The assessment of the genotype’s stability was carried out by the value of the plant breeding
index using the methods of Hangildin V.V. [
          <xref ref-type="bibr" rid="ref11">11</xref>
          ], Francis T.R. and Kannenberg L.W. [
          <xref ref-type="bibr" rid="ref12">12</xref>
          ],
Eberhart S.A. and Russell W.A. [
          <xref ref-type="bibr" rid="ref13">13</xref>
          ], Nassar R. and Huehn M. [
          <xref ref-type="bibr" rid="ref14">14</xref>
          ]. The value of the standard deviation
( 2) and the coeficient of variation ( ) and homeostaticity () of the yield characterize the
absolute and the relative variability of the productivity of the -th cultivar. The regression
coeficient (  ) measures the linear response of the -th variety to varying environments. The
variance of deviation from the regression (2) is a measure of the instability. Mathematical
data processing was performed using the early self-developed computer program “Stability
parameters of crops” [
          <xref ref-type="bibr" rid="ref15">15</xref>
          ].
        </p>
      </sec>
    </sec>
    <sec id="sec-3">
      <title>3. Results and discussion</title>
      <p>To obtain highly productive samples of spring triticale, the patterns of growth and development
of plants in certain climatic conditions, as well as the efect of limiting factors on plants, were
identified. Studies have shown that the sowing time has a significant impact on all traits. Based
on long-term data, pairwise correlations of 10 traits were calculated: stem length, ear length,
number of spikelets per ear, density of an ear, number of grains per ear, grains weight per ear,
weight of 1000 grains, number of stems per plot, yield, number of days between germination
and heading. Analysis of the correlation matrices allowed us to conclude that the traits “spike
length”, “number of grains per ear”, “number of spikelets per ear”, “grains weight per ear” and
“number of days between germination and heading” form a correlation group for both wheat
and for triticale. There was a high level of correlations between all pairs of traits in this group
(0.46 &lt;  &lt; 0.91). The medium linear dependence was observed between the traits “weight of
1000 grain” and “ear density” in wheat ( = 0.31) and between the traits “weight of 1000 grains”
and “stem length” in triticale ( = 0.5). The yield showed a high correlation with the trait
“number of stems per plot” for both crops ( = 0.68;  = 0.71). For a visual representation of
the correlation groups, a cluster analysis was performed based on the correlation matrix of
similarity. As a result, three clusters of traits were identified for wheat and triticale, but the set
of traits was diferent (Figure 1).</p>
      <p>Research data show that at diferent sowing times, the critical stages of plant development
take place in diferent air and soil conditions. This changes the metamerism of the plant and the
ear and afects the productivity of the plant, which is based on numerous correlations between
traits. The lowest yield of all studied genotypes was observed for the third sowing date. The
triticale lines Ukro and Kissa had the highest yield in the trial for the first two sowing dates —
54.2 c/ha and 54.5 c/ha, respectively. They realized their productivity potential mainly due to
the greater number of grains and grains weight from the ear (Figuer 2).</p>
      <p>Summarizing the research results, we can conclude that the plant productivity was most
influenced by the length of the ear, the number of grains in the ear, the number of spikelets in
the ear, the weight of the grains of the ear, and the number of stems from the plot. To a lesser
extent the yield associated with plant height and the weight of 1000 grains. It should also be
noted that the variability of the vegetation conditions of plants in diferent years significantly
influenced the yield. The diference between the average value of the yield for diferent sowing
dates was: 2018 — for wheat 36.3 c/ha, for triticale 76.7 c/ha; 2019 — 31.4 c/ha and 48.4 c/ha;
2020 — 46.0 and 61.5 c/ha, respectively. The maximum yield of the varieties, taking into account
the sowing date, varied from year to year. It was found that under favorable conditions and
correct cultivation techniques, the studied triticale samples are capable of surpassing wheat in
yield (Figure 2).</p>
      <p>The contrasting environmental conditions during the research period and the application of
an integrated approach to the assessment of varieties based on the methods of the static and
dynamic concept made it possible to evaluate the studied genotypes in terms of stability and
adaptability.</p>
      <p>a
b</p>
      <p>The values of the coeficients of variation ( ) and homeostaticity () characterized the
triticale line Sears 57× Ukro as the most stable. The least stable wheat variety was Novosibirskaya 31
(Table 2).</p>
      <p>
        The regression approach to the assessment of stability parameters in accordance with the
method of Eberhart and Russell [
        <xref ref-type="bibr" rid="ref13">13</xref>
        ] showed that varieties Sibirskaya 12 ( = 0.97) and Ukro
( = 1.08), with regression coeficients closest to one, have a medium response to changes in
environmental conditions. This means that a change in the productivity of a variety corresponds
to a change in environmental conditions: in favorable environment it increases moderately, in
unfavorable it decreases moderately.
      </p>
      <p>Varieties Novosibirskaya 31 and Kissa, with a regression coeficient  &gt; 1, are more
responsive to changes in growing conditions and require a higher level of agricultural technology. In
unfavorable environments their productivity sharply decreases and there is a need to replace
8.86
6.81
15.90
13.96
42.55
15.34
10.41

0.68
1.35
0.97
0.48
0.79
1.08
1.66</p>
      <p>
        2
0.0009
0.0004
0.0012
0.0029
0.0002
0.0034
0.0010
2
3.36
3.61
3.50
8.44
1.61
4.44
6.36
them with more adaptive varieties. Varieties Novosibirskaya 15, K-3992 and Sears57× Ukro
with a regression coeficient  &lt; 1 can be recommended for cultivation in unfavorable
environments. The lowest value of variance of deviation from the regression (2) characterizes
variety Sears57× Ukro as the most stable. The non-parametric parameter 2 based on relative
data rankings and proposed by Nassar and Huehn [
        <xref ref-type="bibr" rid="ref14">14</xref>
        ] gave a similar assessment of stability.
In general, all studied varieties showed a high ability to develop under various environmental
conditions.
      </p>
      <p>Based on the evaluation of the plant breeding index (formulas (1), (2)), using data of long-term
breeding trials and self-developed computer software, the following optimal valuable agronomic
traits of hexaploid spring triticale (model of the ideal variety) were determined for the conditions
of the West Siberian forest-steppe zone of the Ob region: plant height — 90 cm; the number
of spikelets per ear — 25 pcs.; the number of grains of the main spike — 65 pcs.; ear length —
12–13 cm; grains weight per spike — 4 g; weight of 1000 grains — 60 g; the number of stems
before harvesting — 167 pcs.; yield — 69.3 c/ha; germination-heading period — 33 days; awnless
or semi-awned forms are acceptable; drought resistance — high; resistance to major diseases
(powdery mildew, brown and stem rusts, loose smut) — high; ecological plasticity is high.</p>
      <p>The productivity potential of triticale significantly exceeds the potential of wheat, but it is
not fully realized.</p>
      <p>Varieties Ukro and Kissa according to the results of cluster analysis are identical to the model
of the ideal variety. Figure 3 graphically presents the elements of the model of an environmental
adaptive variety of spring triticale in comparison with the variety Ukro.</p>
      <p>The analysis of the variability and correlations between valuable agronomic traits of genotypes
indicates the possibility of purposeful breeding for adaptability to the conditions of the region
when creating new high-yielding varieties of spring triticale.</p>
    </sec>
    <sec id="sec-4">
      <title>Acknowledgments</title>
      <p>This work was supported by SFSCAT budget project No. 0778-2020-0001.</p>
    </sec>
  </body>
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