Marker-assisted breeding of a durum wheat

Turkish Journal of Agriculture and Forestry
http://journals.tubitak.gov.tr/agriculture/
Research Article
Turk J Agric For
(2013) 37: 527-533
© TÜBİTAK
doi:10.3906/tar-1207-75
Marker-assisted breeding of a durum wheat cultivar for γ-gliadin and
LMW-glutenin proteins affecting pasta quality
1,
1
2
Ahmet YILDIRIM *, Özlem ATEŞ SÖNMEZOĞLU , Abdulvahit SAYASLAN ,
2
1
3
Mehmet KOYUNCU , Tuğba GÜLEÇ , Nejdet KANDEMİR
1
Department of Biology, Karamanoğlu Mehmetbey University, Karaman, Turkey
2
Department of Food Engineering, Karamanoğlu Mehmetbey University, Karaman, Turkey
3
Department of Field Crops, Gaziosmanpaşa University, Tokat, Turkey
Received: 03.08.2012
Accepted: 14.12.2012
Published Online: 28.08.2013
Printed: 25.09.2013
Abstract: The pasta quality of durum wheat is one of the most important properties for the industry and consumers. Therefore, breeding
for improved grain quality without yield penalties, using modern breeding methods, has been a primary objective in durum wheat
breeding programs in recent years. In this study, 2 important gene regions that encode pasta-quality associated proteins (γ-gliadin 45
and LMW-2 glutenin) were transferred to a registered Turkish durum wheat variety, Sarıçanak-98, from a high-quality Canadian durum
wheat cultivar, Kyle, through a marker-assisted backcross breeding method. Each of the F1 and backcross (BC) plants were backcrossed
4 times to the recurrent parent, and the backcrossed plants carrying the targeted gene regions in all generations were selected by markerassisted selection (MAS). DNA markers in combination with A-PAGE were used for tracking the introgression of the targeted gene
regions. Transfer of Gli-B1 locus encoding γ-gliadin 45 and Glu-B3 locus encoding LMW-2 glutenin to the wheat variety Sarıçanak-98
led to a considerable increase in protein content and gluten quality.
Key words: Durum wheat, Triticum durum, γ-gliadin 45, LMW-2 glutenin, backcross breeding, MAS, SSR
1. Introduction
Wheat quality is simply defined as its suitability for a
given final product. For durum wheat, quality means
its suitability for pasta processing. The quality of pasta
products is strongly correlated with the physical and
chemical properties of durum wheat. Kernel vitreousness,
protein content and quality (proper gluten strength),
milling properties (semolina yield and ash content), yellow
pigment content, and activities of lipoxygenase (LOX),
peroxidase (POD), and polyphenol oxidase (PPO) enzymes
are among the well-documented quality parameters
(Clarke et al. 1998; Borrelli et al. 1999; Troccoli et al. 2000;
Morris 2004). Of these quality parameters, protein content
and quality, yellow pigment content, and the activities of
the oxidative enzymes are of vital importance for pasta
quality, as they overwhelmingly determine the so-called
al dente cooking characteristics and bright yellow color of
pasta products (Troccoli et al. 2000; Aalami et al. 2007).
However, protein content and gluten quality (including
gliadin and glutenin proteins) are the 2 most important
variables in determining pasta cooking properties (D’Edigio
et al. 1990; Kovacs et al. 1995; Blanco et al. 1996; Troccoli
et al. 2000). Moreover, there is an important correlation
*Correspondence: [email protected]
among specific gliadin and glutenin proteins and the gluten
strength. The most important of those proteins are γ-gliadin
45 and LMW-2 glutenin, which are linked very tightly to
each other. γ-gliadin 45 is strongly correlated with high
gluten strength and pasta quality (Gupta et al. 1994; Kovacs
et al. 1995; D’Ovidio and Porceddu 1996). It has been
reported that transfer of a Gli-B1 locus encoding γ-gliadin
45 and a Glu-B3 locus encoding LMW-2 glutenin together
into a variety leads to improvement in the pasta cooking
quality of durum wheat (D’Ovidio et al. 1992; Kovacs et al.
1994; Clarke et al. 1998).
Turkey is blessed with agro-climatic regions suitable for
the cultivation of high-quality durum wheat. It is among the
top 3 durum wheat producing countries in the world with 3
Mt of production during 2009. However, only about 30%–
40% of the total durum wheat is able to meet the quality
criteria required by the pasta industry (IGC 2003), leading
to a rise in importation of durum wheat with desirable pasta
qualities each year. Therefore, the pasta processing quality
of Turkish durum wheat cultivars must be improved using
modern breeding methods, without adversely affecting
grain yield. For this purpose, existing varieties may be
improved through introgression of genes/QTLs associated
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YILDIRIM et al. / Turk J Agric For
with pasta quality. Sarıçanak-98, a durum wheat cultivar,
is widely grown in southeastern Turkey due to its high
yield potential, resistance to common wheat diseases in the
region, and tolerance of poor soil conditions, particularly
in terms of microelement deficiencies (Özberk et al. 2010;
CRIFC 2011). However, the cultivar Sarıçanak-98 suffers
from poor end-use quality, as it has the γ-gliadin 42 and
LMW-1 proteins, which are related to poor pasta quality.
Sarıçanak-98 may be further improved by transfer of
γ-gliadin 45 and LMW-2 glutenin proteins associated with
superior pasta-cooking quality. The aim of this study was
to transfer 2 important gene regions encoding γ-gliadin
45 and LMW-2 glutenin proteins from the high-quality
Canadian durum wheat cultivar Kyle to Sarıçanak-98
through a marker-assisted backcross breeding method.
2. Materials and methods
2.1. Plant materials
Sarıçanak-98, widely grown in Turkey and lacking the
γ-gliadin 45 and LMW-2 glutenin encoding genes, was
used as the recurrent parent. The donor parent was a highquality Canadian durum wheat cultivar, Kyle. In each
generation, F1 and BC plants were backcrossed 4 times
(BC4) to the recurrent parent, and progenies carrying
the targeted gene regions were selected by MAS. In each
generation, an average of 50–100 BC plants were screened
by molecular DNA markers, and all screening results were
verified and tested with A-PAGE. Advanced breeding lines
(ABLs) were obtained by selfing BC4F2 plants twice. Seeds
of the lines were multiplied by growing in field conditions
during the 2009–2010 growing seasons.
The cultivars Marquis and Kyle were used as check
genotypes for the identification of γ-gliadin 42/45 bands
on A-PAGE.
2.2. DNA screening
Seeds of F1 and backcross plants were cut into halves. One
half was used for A-PAGE and the other half, including
the embryo, was germinated and used for screening with
DNA markers. DNA was extracted from the leaf of each
plant using a genomic DNA purification kit (Fermentas
Life Sciences).
DNA samples were screened. For this purpose, the genebased marker GAG5-6 (Von Büren et al. 2000), linked to
both Gli-B1 and Glu-B3 loci, was used for screening of the
F1 and backcross population for the presence of γ-gliadin
45 and for LMW-2 glutenin alleles.
2.3. PCR amplification
Polymerase chain reactions (PCRs) were performed under
the conditions given elsewhere (Von Büren et al. 2000)
with some modifications. PCR reaction volume was 40
µL. PCR reactions contained 50–100 ng of genomic DNA,
0.25 µM of each primer, 0.2 µM dNTP mix, 2.5 µM MgCl2,
10X PCR buffer, and 0.5 units of Taq DNA polymerase.
528
The PCR was run on a Thermo (Px2) thermal cycler as
follows: an initial denaturation step of 3 min at 94 °C was
followed by 32 cycles of denaturation for 1 min at 94 °C,
annealing for 1 min at 60 °C, extension for 1 min at 72 °C,
and conclusion with a final extension step for 5 min at 72
°C. PCR products were analyzed on 3% metaphore agarose
gels or 1% agarose gels. Electrophoresis was conducted at
90 W of constant power for 3–4 h.
2.4. A-PAGE
The gliadins (γ-gliadin 42/45) of wheats were identified
using the A-PAGE method that was originally described
by Bushuk and Zillman (1978) and modified by Khan et al.
(1985). In each generation, an average of 50–100 BC plants
were screened by A-PAGE.
2.5. Embryo culture
To speed up BC generations, some BC plants were
generated by immature embryo cultures following the
process described by Arzani and Mirodjagh (1999).
2.6. Quality measurements on grain
Selected grain properties associated with quality were
measured using the following methods in 3 replications:
kernel vitreousness by Köksel et al. (2000), thousand
kernel weight and kernel size distribution by Elgün et
al. (2002), and test weight, kernel color, yellow pigment
content, SDS sedimentation volume, moisture content,
protein content, and ash content by the AACC methods
of 55-10, 14-22, 14-50, 56-70, 44-15A, 46-10, and 08-01,
respectively (AACC 2000).
Lipoxygenase (LOX) activity was determined by
spectroscopic measurement of a conjugated diene
formation upon the reaction of wheat extracts with a
linoleic acid substrate, as described by Rani et al. (2001)
and Aalami et al. (2007). One unit of LOX activity (EU)
was described as 1.0 unit min–1 change in the absorbance
under the assay conditions and reported as EU per g
wheat. Polyphenol oxidase (PPO) activity was determined
following the method of Coseteng and Lee (1987), as
modified by Aalami et al. (2007), where color changes
in the mixture of wheat extract–catechol substrate were
monitored at 420 nm. One unit of PPO activity was
described as 0.1 unit min–1 change in the absorbance under
the assay conditions and reported as EU per g wheat.
Peroxidase (POD) activities were measured according to
the method of Aparicio-Cuesta et al. (1992), modified by
Aalami et al. (2007), where colored product formation
upon the reaction of wheat extracts with an o-dianisidin
substrate in the presence of hydrogen peroxide, was
monitored at 460 nm. One unit of POD activity was
described as 1.0 unit min–1 change in the absorbance
under the assay conditions and reported as EU/g wheat.
2.7. Quality measurements of semolina
Selected quality characteristics of semolina were measured
in 3 replications as follows: a dark-colored speck count
YILDIRIM et al. / Turk J Agric For
alleles through molecular markers (Figure 1) and A-PAGE
(Figure 2) in each generation.
A 1-to-1 segregation ratio was clearly seen in the
BC1F1 generation. Fifty percent of the backcross plants
carried the recurrent parent allele (Sarıçanak-98), while
the rest were heterozygous, as expected.
In each backcross generation, screening was carried out
in combination with molecular DNA markers and A-PAGE.
As a result of the screening, heterozygous backcross plants
having the targeted alleles were determined and the BC4F1
seeds were obtained through hybridization of those plants.
The BC4F2 seeds were obtained by selfing of BC4F1 plants.
In the BC4F2 plants, homozygous plants (25%) were
identified by molecular (Figure 3a) and A-PAGE screening
(Figure 3b), and homozygous BC4F2 seeds were increased
by selfing to use for the yield trials later.
based on Elgün et al. (2002) and milling yield, color,
pigment content, gluten content/index, moisture content,
protein content, and ash content according to the AACC
methods of 26-41/42, 14-22, 14-50, 38-12A, 44-15A, 4610, and 08-01, respectively (AACC 2000).
3. Results
3.1. Molecular and biochemical marker-assisted
selection
The genes encoding γ-gliadin 45 and LMW-2 glutenin
proteins, which affect end-use quality of durum wheat,
were transferred to the Turkish durum wheat cultivar
Sarıçanak-98 following a marker-assisted backcross
breeding program. The F1 and BC plants were backcrossed
4 times to the recurrent parent, and the backcross plants
(from BC1F1 to BC4F1) were screened for the targeted
K
S
-45
-42
K
S
-45
-42
H
H
A
H
A
A
H
H
A
A
A
A
H
H
H
A
H
B
B
B
H
H
H
Figure 1. Representative PCR profiles of parents and BC1F1 plants due to GAG 5-6
markers linked to Gli-B1 locus. (H: plants heterozygous for targeted gene regions.)
M
K
S
1
2
3
4
5
6
7
8
9
10 M
A
H
A
H
A
H
H
A
H
H
-42
-45
Figure 2. Representative profiles of parents and BC1F1 plants screened with
A-PAGE. (A: BC1F1 plants homozygous for the recurrent parent alleles, H:
BC1F1 plants heterozygous for the targeted gene regions, K: cv. Kyle, S: cv.
Sarıçanak-98).
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YILDIRIM et al. / Turk J Agric For
a
b
K
S
H
H
B
A
H
H
1
B
2
K S
M
H H -45 -42
K
S
A
A
H
A
A
H
A
H
H
H
3
4
5
6
7
B
H
A
B A
8
9
10 11 12
B
H H
H H
H
Figure 3. Representative profiles of parents and BC4F2 plants screened with GAG 5-6 markers (a) and A-PAGE (b). (A:
Backcross plants homozygous for the recurrent parent alleles, B: Backcross plants homozygous for the donor parent alleles, H:
Heterozygous backcross plants.)
Immature embryo cultures were used to reduce the
time required for seed generation after hybridization
and acceleration of the backcross processes. Half of the
BC seeds in each generation were germinated through
immature embryo cultures and the rest were left on BC
spikes for normal maturation. Thus, each generation was
produced approximately 40 days earlier. Consequently,
more than 2 generations were obtained in a year with the
help of the embryo culture approach.
3.2. Quality improvement
Selected physical, chemical, and technological properties
of the advanced breeding line (BC4F4) and its parents
(cultivars Kyle and Sarıçanak-98) were investigated and
the findings summarized in Tables 1–3. The breeding
line and its parents were fairly comparable in terms of
kernel physical properties, although the differences were
statistically significant (P < 0.05) (Table 1). However,
the breeding line significantly differed from its parents
with respect to gluten quality and protein content (Table
2). Gluten quality of the breeding line was significantly
improved, as indicated by its higher SDS sedimentation
and specific sedimentation volumes, which was the aim
of this work. Additionally, protein content of the breeding
line was significantly higher (P < 0.05) than that of its
recurrent parent, Sarıçanak-98. Indeed, simultaneous
improvement of gluten quality and protein content is a
significant achievement for the advanced breeding line.
Initial observations indicated that yield of the ABL did not
Table 1. Physical properties of the advanced breeding line and its parents.a,b
Parents /Advanced
breeding line
Kyle
Sarıçanak-98
ABL (BC4F4)
Thousand kernel
weight (g)
Test weight
(kg hL–1)
36.7 a
36.8 a
35.5 b
79.2 c
82.3 a
80.6 b
Kernel vitreousness
(%)
>2.8 mm
99.8 a
99.1 b
99.5 ab
23.6
27.4
47.3
Kernel size distribution (%)
>2.5 mm
>2.2 mm
>2.0 mm
Homogeneity
37.3
47.6
34.0
29.1
18.6
14.6
10.2
6.5
4.1
Heterogeneous
Homogeneous
Homogeneous
14% moisture basis bDifferent letters in a column indicate significant difference (P < 0.05)
ABL: Advanced breeding line, BC: Backcross
a
Table 2. Chemical and technological properties of the advanced breeding line and its parents.a,b
Parents / Advanced
breeding line
Kyle
Sarıçanak-98
ABL (BC4F4)
Moisture
content (%)
Ash content
(%)
Protein content
(%)
Sedimentation
volume (mL)
Specific sedimentation
volume (mL)
10.4 ns
9.8 ns
10.3 ns
1.54 b
1.48 b
1.65 a
14.8 a
13.8 b
14.8 a
22.1 a
17.9 b
22.5 a
1.49 a
1.30 b
1.52 a
a
14% moisture basis bDifferent letters in a column indicate significant difference (P < 0.05) ns: Not significant
ABL: Advanced breeding line, BC: Backcross
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YILDIRIM et al. / Turk J Agric For
show any difference from Sarıçanak-98. However, reliable
results can be obtained from replicated field trials in the
wheat’s natural environment .
As seen in Table 3, yellow pigment content and color
properties (L* and b*) of the breeding line were significantly
higher (P < 0.05) than those of the recurrent parent. On
the other hand, the breeding line and its recurrent parent
were statistically comparable in terms of their oxidative
enzymes (LOX, PPO, and POD) that contribute to the
bright yellow color of pasta products.
Semolina yields of the breeding line and its parents
were quite similar; however, the breeding line had
significantly higher levels of pigment and protein with
slightly improved gluten quality (Table 4). It is obvious
that quality measurements recorded on the grain (Tables 2
and 3) were in close agreement with those recorded on the
semolina (Table 4).
4. Discussion
Pasta products are appreciated worldwide as an end-product
of durum wheat. Considering increased competition in
the global pasta market, improving the end-use quality of
durum wheat is essential. It has been reported that there is a
positive relationship between γ-gliadin 45 protein (LMW2 glutenin) and the optimum gluten strength associated
with the al dente cooking quality of pasta (Kovacs et al.
1995; Troccoli et al. 2000). It has been widely reported that
the gluten quality of durum wheat containing γ-gliadin
45 and LMW-2 glutenin is better than that of γ-gliadin 42
and LMW-1 glutenin (Damidaux et al. 1978; Chihab 1990;
Kaan et al. 1995; Nachit et al. 1995; Pena 2000; Edwards
et al. 2007; Yüksel 2009). It is also well known that gluten
quality and the protein content of the durum are the main
parameters that determine pasta quality (Bushuk 1998;
Troccoli et al. 2000).
During this study, important gene regions encoding
pasta-quality associated proteins (γ-gliadin 45 and LMW2 glutenin) were transferred from the cultivar Kyle to
the cultivar Sarıçanak-98, leading to the development
of an advanced breeding line with improved quality. As
expected, this line has increased gluten quality, as judged
by the sedimentation and gluten index tests. Additionally,
the protein content of the breeding line also increased.
Besides the protein content and gluten quality, a yellow
pigment color of the grain also contributes to improved
pasta quality. Pigment contents ranging from 4 to 8 mg
kg–1 in durum wheat have been reported by different
researchers. The pigment content (7.55 mg kg–1) of the
breeding line developed during the present study was
higher than the pigment content (4 to 8 mg kg–1) reported
in earlier durum wheat studies (Kaan et al. 1995; Köksel et
al. 2000; Troccoli et al. 2000; Sakin et al. 2011). Thus, the
breeding lines developed should produce yellow-colored
pasta products, preferred by consumers.
The results of this study confirm that marker-assisted
selection and backcross breeding can be successfully
Table 3. Color properties and oxidative enzyme activities of the advanced breeding line and its parents.a,b
Parents / Advanced
breeding line
Kyle
Sarıçanak-98
ABL (BC4F4)
Pigment
Content
(mg/kg)
L*
a*
7.58 a
6.41 b
7.55 a
77.0 a
59.3 b
76.5 a
5.05 b
5.22 a
5.17 a
b*
LOX
Activity
(EU g–1)
POD
Activity
(EU g–1)
PPO
Activity
(EU g–1)
18.59 a
17.37 b
18.59 a
44.9 b
48.1 a
48.5 a
29.3 b
62.7 ab
78.9 a
6.8 ns
6.9 ns
11.1 ns
Color
14% moisture basis bDifferent letters in a column indicate significant difference (P < 0.05) ns: not significant
ABL: Advanced breeding line, BC: Backcross
a
Table 4. Chemical and technological properties of semolina milled from the advanced breeding line and its parents.a,b
Parents /
Advanced
breeding line
Semolina
yield
(%)
Pigment
content
(mg kg–1)
Speck count
(number 100
cm–2)
Ash
content
(%)
Protein
content
(%)
Wet gluten
content
(%)
Dry gluten
content
(%)
Gluten
index
Kyle
Sarıçanak-98
ABL (BC4F4)
54.7
55.4
54.1
7.31 a
6.00 c
6.85 b
110
110
75
0.59 a
0.62 a
0.55 b
13.8 a
12.9 b
13.8 a
44.0 ns
38.2 ns
39.4 ns
16.5 ns
14.6 ns
15.7 ns
80 b
91 ab
98 a
14% moisture basis bDifferent letters in a column indicate significant difference (P < 0.05) ns: not significant
ABL: Advanced breeding line, BC: Backcross
a
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YILDIRIM et al. / Turk J Agric For
employed in wheat breeding programs and that molecular
markers can be used alone or in combination with the
A-PAGE technique in each backcross generation. Qualitytargeted breeding through marker-assisted backcross
breeding in this work was completed in about 3 years by
raising 3 generations per year. With the help of markerassisted selection, breeding time was reduced and the
efficiency of backcross breeding was increased, leading to
the development of a durum wheat candidate with elevated
protein content and gluten quality.
Acknowledgments
This study was supported by the Scientific and
Technological Research Council of Turkey (TÜBİTAK;
Project No. 107O004) within the framework of COSTFA-0604 TritiGen Action.
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