Received: 27 March 2017
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Accepted: 27 July 2017
DOI: 10.1111/pbr.12523
ORIGINAL ARTICLE
Molecular cytogenetic characterization and phenotypic evaluation of new wheat–rye lines derived from hexaploid triticale ‘Certa’ 3 common wheat hybrids Caiyun Liu1,2*
| Quanhao Song1* | Hongjun Zhang3 | Zujun Yang3 | Yin-Gang Hu1,4
1 State Key Laboratory of Crop Stress Biology for Arid Areas, College of Agronomy, Northwest A&F University, Yangling, Shaanxi, China 2
Dezhou Academy of Agricultural Sciences, Dezhou, Shandong, China 3
School of Life Science and Technology, University of Electronic Science and Technology of China, Chengdu, Sichuan, China 4 Institute of Water Saving Agriculture in Arid Regions of China, Northwest A&F University, Yangling, Shaanxi, China
Correspondence Yin-Gang Hu, State Key Laboratory of Crop Stress Biology for Arid Areas, College of Agronomy, Northwest A&F University, Yangling, Shaanxi, China. Email:
[email protected] Funding information Sub-project of the 863 Program of the Ministry of Science and Technology, Grant/ Award Number: 2013AA102902; China 111 Project of the Ministry of Education of China, Grant/Award Number: B12007; Ministry of Education of China; Science and Technology Innovation Team Plan of Shaanxi Province, Grant/Award Number: 2014KCT25; the Open Project of the State Key Laboratory of Crop Stress Biology for Arid Areas, Grant/Award Number: CSBAA2017006
Abstract Hexaploid triticale contains valuable genes from both tetraploid wheat and rye and plays an important role in wheat breeding programmes. In order to explore the potential of hexaploid triticale ‘Certa’ in wheat improvement, two crosses were made using ‘Certa’ as female parent, and common wheat cultivars ‘Jinmai47’ (JM47) and ‘Xinong389’ (XN389) as male parents. The karyotyping of BCF4:5 lines from Certa/JM47//JM47 and F5:6 lines from Certa/XN389 was investigated using sequential fluorescence in situ hybridization (FISH). One 1B(1R) substitution line and five 1BL.1RS whole-arm translocation lines were identified, one of which was found lacking x-secalin locus. Many structural alterations on wheat chromosomes were detected in the progeny. Great morphologic differences resulting from genetic variations were observed, among which the photosynthetic capability was increased while grain quality was slightly improved. Compared with both parents, the stripe rust resistance at adult stage was increased in lines derived from Certa/JM47// JM47, while it was decreased in lines derived from Certa/XN389. These newly developed lines might have the potential to be utilized in wheat improvement programmes. KEYWORDS
chromosome translocation, fluorescence in situ hybridization, phenotypic traits, stripe rust resistance, Triticum aestivum
Communicated by: T. Miedaner
1 | INTRODUCTION
diversity of common wheat (Reif et al., 2005; Warburton et al., 2006). The wild relatives of wheat possess abundant desirable traits
Wheat (Triticum aestivum L.) is a major cereal crop and plays an
for wheat improvement (Colmer, Flowers, & Munns, 2006; Friebe,
important role in ensuring food security worldwide. However, with
Jiang, Raupp, McIntosh, & Gill, 1996). Distant hybridization and chro-
global climate change, the stability and productivity of wheat are
mosome engineering make it possible to transfer useful traits from
affected seriously by various biotic and abiotic stresses. Meanwhile,
wild relatives into common wheat, and numerous germplasm with
modern selective breeding procedures have narrowed the genetic
alien chromosomes have been created. In most cases, those germplasm are used as genetic tools, yet little is known regarding their
*These authors contributed equally to this work.
Plant Breeding. 2017;1–11.
potential in breeding programmes. wileyonlinelibrary.com/journal/pbr
© 2017 Blackwell Verlag GmbH
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ET AL.
Hexaploid triticale (X Triticosecale Wittmack, 2n = 69 = 42,
materials, probe labelling and in situ hybridization were performed as
AABBRR), derived from crossing durum wheat (Triticum turgidum
described by Kato, Lamb, and Birchler (2004) and Han, Lamb, and
subsp. durum (Desf.) Husn., 2n = 49 = 28, AABB) with rye (Secale
Birchler (2006).
cereale L., 2n = 29 = 14, RR), consists of all the chromosomes of
Oligonucleotide probes used in this study were synthesized by
wheat A and B genomes and rye R genome. It is a desirable germ-
Shanghai Invitrogen Biotechnology Co. Ltd. (Shanghai, China). Oligo-
plasm due to the high biomass and grain yield with the growth vig-
pSc200 and Oligo pSc119.2-1 were 50 end-labelled with 6-carboxy-
our and environmental tolerance from rye (Kang et al., 2016).
fluorescein (6-FAM), and Oligo pTa535-1 was 50 end-labelled with 6-
Triticale materials have been used to produce wheat–rye chromo-
carboxytetramethylrhodamine (Tamra) (Tang, Yang, & Fu, 2014).
some addition, substitution and translocation lines. Desirable traits
Images were taken using an epifluorescence microscope (BX51,
or genes with disease resistances and end-use quality have been
Olympus) equipped with a cooled charge-coupled device camera
successfully transferred into common wheat from hexaploid triticales
operated with HCIMAGE Live software (version 2.0.1.5) and pro-
(Yang et al., 2006). For instance, several hexaploid triticales with the
cessed with Photoshop CC 2014.
rye genome from Secale strictum (J. Presl) J. Presl exhibited novel strong resistance to Russian wheat aphid (Nkongolo & Comeau, 1998), and similar studies were carried out on wheat–S. montanum
2.3 | Rye-specific molecular marker analysis
(Secale montanum Guss.) addition, substitution and translocation lines
Fifty-five pairs of rye-specific PCR-based Landmark Unique Gene
(Montero, Sanz, & Jouve, 1986). In the process of transferring rye
(PLUG) markers assigned to rye chromosomes (Table S1) were used
chromatin into wheat using triticale as the donor, recombination
to detect the rye fragments, and the primers were synthesized
could occur between the A and B genome’ chromosomes of triticale
according to Ishikawa et al. (2009) and Li et al. (2013). A molecular
and common wheat; therefore, the A and B genome’ chromosomes
marker for rye-specific x-secalin (SEC-1b) was used as described by
could be altered accompanying with allopolyploidization in their pro-
Zhang et al. (2002). The PCR amplification, enzyme digestion and
geny (Levy & Feldman, 2004). In addition, previous studies showed
electrophoresis were carried out as described by Li et al. (2013).
that rye chromosomes added to a wheat background could induce modifications in wheat chromosomes, which may represent new genetic changes and epigenetic variations (Tang, Li, et al., 2014). Hexaploid triticale ‘Certa’ is a spring triticale cultivar developed
2.4 | Agronomic traits assessment and stripe rust resistance evaluation
from the cross of two triticales (Hare 263 and Civet) at CIMMYT
The field experiment was conducted during the growth season of
(International Maize and Wheat Improvement Center) in 1989, with a
October 2014 to June 2015 in the farm of the Northwest A&F
significant improvement in test weight and Hagberg falling number,
University, Yangling, Shaanxi, China. Wheat lines were arranged in a
and displayed moderate resistant to the prevalent races of stem rust
randomized complete block design with three replications.
and leaf rust, highly resistant to common bunt, and resistant to com-
The time of heading (Z55, Stapper, 2007) and flowering (Z65)
mon root rot (McLeod, Pfeiffer, DePauw, & Clarke, 1996). In the pre-
was visually recorded when 50% of the plants per plot had reached
sent study, two crosses were made using ‘Certa’ as the female parent,
the stage. Days to heading (DH) and days to flowering (DF) from
and common wheat cultivars ‘Jinmai47’ (JM47) and ‘Xinong389’
seedling emergence stage (Z10) were calculated. The length and
(XN389) as male parents. The chromosome constitutions of the BCF4:5
width of flag leaf were measured at early grain-filling stage (Z70).
and F5:6 lines derived from Certa/JM47//JM47 and Certa/XN389,
Plant height (PH), spike length (SL), tiller number (TN), spikelet num-
respectively, were characterized, and their agronomic traits, photosyn-
ber (SN) and grain number per spike (GNS) were determined at
thetic capability, grain quality and resistance to stripe rust were evalu-
maturity stage (Z92). At harvest, 10 individual plants of each replica-
ated for their potential utilization in wheat breeding programmes.
tion were hand-cut at ground level. Samples were dried at 35°C for 3 days and weighed before and after threshing as the above-ground
2 | MATERIALS AND METHODS 2.1 | Plant materials
biomass and grain yield. After that, biomass per plant (BP), grain yield per plant (YP) and harvest index (HI) were calculated. One thousand grains of each genotype were weighed and the values were averaged as TGW. Wheat lines were inoculated with races of
The plant materials used in this study included hexaploid triticale ‘Certa’
Puccinia striiformis f. sp. tritici (CYR31 and CYR32) and evaluated for
(2n = 69 = 42, AABBRR), common wheat JM47 and XN389 (2n = 69
adult stage resistance to stripe rust in the field. The infection type
= 42, AABBDD), BCF4:5 lines derived from recurrent backcross of ‘Certa’
(IT) was recorded based on the 0 to 4 scale according to Bariana
with JM47 and F5:6 lines derived from ‘Certa’ 9 XN389 (Figure S1).
and McIntosh (1993).
2.2 | Sequential fluorescence in situ hybridization
2.5 | Photosynthesis characteristics assessment
Fluorescence in situ hybridization (FISH) was used to analyse the
Relative chlorophyll content (SPAD value) of flag leaves was mea-
mitotic metaphase cells of wheat lines. Chromosome spreads of
sured at heading (Z55), flowering (Z65) and early grain-filling (Z70)
LIU
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3
stages with a SPAD-502 hand-held chlorophyll meter (Minolta Cam-
pSc200 as a probe, indicating that there was no rye chromosome
era Co. Ltd., Japan).
detected. Among the eight lines from Certa/XN389, strong signals
Photosynthesis characteristics including net photosynthetic rate
were observed in six lines, which contained one pair of hybridized
(Pn), stomatal conductance (Sc), intercellular CO2 concentration (Ci)
chromosomes using Oligo-pSc200 as a probe (Figure 2). Strong sig-
and transpiration rate (Tr) were measured on flag leaves at heading
nals appeared at the telomeric regions of both long and short arms
(Z55), flowering (Z65) and early grain-filling (Z70) stages using a LI-
in line 12-1-2 (Figure 2a, b), while strong signals were only observed
6400XT portable photosynthesis system (Li-Cor Inc., Lincoln NE,
at the telomeric regions of short arms in the other five lines (lines
USA). The measurements were conducted under 400 lmol CO2
12-4-1, 12-4-2, 12-6-2, 12-10-1 and 12-11-4). The subsequent FISH
mol1 air concentration and 1000 lmol m2 s1 photosynthetic
analysis using Oligo-pSc119.2-1 and Oligo-pTa535-1 as probes
photon flux density.
revealed that line 12-1-2 was an 1B(1R) substitution line, and the other five lines were 1BL.1RS translocation lines (Figure 2c, d). Additionally, differences in the FISH signal patterns of wheat
2.6 | Grain quality analysis
chromosomes between the derived lines and their parents suggest
The protein, starch, fibre and gluten content, hardness, Zeleny sedi-
that alterations of wheat chromosomes occurred. For instance,
mentation and SDS sedimentation values of whole grains were
among lines derived from Certa/XN389, line 12-1-2 gained Oligo-
determined using the near-infrared spectrometer DA7200 (Perten,
pSc119.2-1 signal at the terminal region of 1DS arm, lost Oligo-
Sweden) with a model for wheat grain analysis. The high molecular
pSc119.2-1 and enhanced Oligo-pTa-535-1 signal at the terminal
weight glutenin subunits (HMW-GS) were analysed by the sodium
region of 3DS arm (Figure 3c). The Oligo-pSc119.2-1 signal loss
dodecyl sulphate–polyacrylamide gel electrophoresis (SDS-PAGE)
appeared at the terminal regions of chromosomes 2B, 6B or 7B in
method.
the other lines (Figure 3).
3 | RESULTS
3.2 | Detection with rye-specific PLUG markers
3.1 | Chromosome composition by sequential FISH analysis
Wheat lines were further analysed with 55 pairs of rye-specific PLUG primers assigned to 1R to 7R and rye-specific x-secalin marker SEC-1b (Table S1). Rye-specific bands were amplified by PLUG
Oligo-pSc200 produced hybridization signals on all 14 rye chromo-
markers TNAC1009, TNAC1019 and TNAC1063 assigned to 1RS in
somes in hexaploid triticale ‘Certa’, but no signals were observed on
the 1B(1R) substitution line 12-1-2, and the translocation lines 12-4-
wheat chromosomes (Figure 1a), which indicated that Oligo-pSc200
1, 12-4-2, 12-6-2, 12-10-1 and 12-11-4 (Figure 4), which verified
allowed rye chromosomes to be unambiguously distinguished from
the results of FISH analysis. Meanwhile, rye-specific x-secalin bands
wheat chromosomes. The combination of Oligo-pSc119.2-1 and
were not detected in line 12-6-2, suggesting that it was a x-secalin
Oligo-pTa535-1 probes could distinguish the individual chromosomes
gene deletion line. Except the substitution and translocation lines, no
of the wheat A and B genomes as well as the entire rye genome
rye-specific band was amplified in the other lines (Figure S2).
(Figure 1b). The chromosome composition of the lines derived from ‘Certa’ and common wheat was demonstrated by sequential FISH analysis.
3.3 | Agronomic traits and stripe rust resistance
The somatic chromosome number of the six lines from Certa/
Compared with JM47 and XN389, ‘Certa’ was superior in PH, SL,
JM47//JM47 was 42, and no signal was observed with Oligo-
SN, GNS, BP and YP (Table 1). ‘Certa’ was 3–5 days earlier than
(a)
F I G U R E 1 FISH analyses using OligopSc200 (a), Oligo-pSc119.2-1 and OligopTa535-1 (b) as probes on root tip metaphase chromosomes of hexaploid triticale ‘Certa’ (a) 40 ,6-diamidino-2phenylindole (DAPI), blue fluorescence; Oligo-pSc200, green fluorescence. (b) Oligo-pTa535-1, red fluorescence; OligopSc119.2, green fluorescence. Bar = 10 lm. Arrows indicate the rye chromosomes
(b)
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(b)
(c)
(d)
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F I G U R E 2 FISH analyses using OligopSc200 (a, c), Oligo-pSc119.2-1 and OligopTa535-1 (b, d) as probes on root tip metaphase chromosomes of lines derived from Certa/XN389 (a, b) Line 12-1-2; (c, d) line 12-11-4. Arrows indicate rye chromosomes. Bar = 10 lm
(a)
(c)
(e)
(b)
(d)
(f)
F I G U R E 3 FISH karyotypes of lines derived from Certa/XN389. (a) ‘Certa’; (b) XN389; (c) line 12-1-2; (d) line 12-2-1; (e) line 12-4-2; (f) line 12-11-4. Wheat chromosomes with changes are indicated in white box. Bar = 10 lm
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FIGURE 4
5
Detection of Sec-1 locus in lines from Certa/XN389. Rye-specific bands are indicated with arrows
common wheat to reach the heading stage, but showed the similar
while the content of protein, gluten and starch of ‘Certa’ was
flowering time as the common wheat varieties.
between JM47 and XN389 (Table 3). Among the lines derived from
Lines derived from Certa/JM47//JM47 did not improve much in
Certa/JM47//JM47, line 7-3-1 expressed significantly higher values
yield components (Figure 5a). Most of the lines showed short flag
than JM47 in all the quality traits except starch content, and line 7-
leaves as ‘Certa’, while line 10-3-1 had increased FLW and much lar-
2-1 showed clearly higher Zeleny sedimentation and SDS sedimenta-
ger FLA. In addition, DH and DF were shortened in most lines com-
tion values than JM47. The lines carrying 1RS derived from Certa/
pared with both parents (Table 1). Lines derived from Certa/XN389
XN389 showed significantly lower protein content, gluten content
benefited significantly from both parents in agronomic traits (Fig-
and Zeleny sedimentation values, while significantly increased starch
ure 5b). The five 1BL.1RS translocation lines advanced DH and DF
content than XN389.
for approximately 3–4 days compared with XN389, while the 1B(1R)
Given the codominance of Glu-1 alleles, each functional allele
substitution line shortened the DH and DF for approximately 11 and
encodes a distinguishable HMW-GS and the HMW secalins of rye
8 days, respectively. Several lines carrying 1RS produced more spike-
consisted of x-type and y-type subunits (De Bustos & Jouve, 2003).
let per spike, with increased YP and BP compared with XN389.
The HMW-GS allelic composition of JM47 was determined as Null,
‘Certa’ was nearly immune to the tested isolates of stripe rust,
1Bx7+1By9 and 1Dx3+1Dy12, while that of XN389 was 1Ax1,
suggesting that ‘Certa’ might carry the resistant genes to stripe rust
1Bx7+1By9 and 1Dx2+1Dy12 (Table 3). The HMW-GS of ‘Certa’
derived from rye. Common wheat variety JM47 was highly suscepti-
consisted of 1Ax1, 1Bx7+1By18 and 1Rx6+1Ry13. All the lines
ble, while the lines derived from ‘Certa’ and JM47 showed moderate
derived from ‘Certa’ and JM47 showed the same pattern as JM47 in
resistance to stripe rust. Carrying the resistance gene Yr26 (Li et al.,
the HMW-GS combination. The 1BL.1RS translocation lines derived
2008) located on 1B chromosome, XN389 was nearly immune to the
from Certa/XN389 had the similar HMW-GS patterns as XN389.
tested isolates, while most lines derived from Certa/XN389 showed
However, in line 12-1-2, the substitution line derived from Certa/
high resistance to stripe rust, but less resistant than both parents,
XN389, 1Ax1 was not detected, but 1Dx2+1Dy12 was detected and
indicating that Yr26 may have not been transferred into the progeny.
1Bx7+1By9 was replaced by 1Rx6+1Ry13. This may suggest that the allele encoding 1Rx6+1Ry13 was located on the long arm of 1R
3.4 | Photosynthetic characteristics
chromosome. Interestingly, a novel pattern 1Dx4+1Dy12 was observed in line 12-6-2 (Figure S3), suggesting the coding sequences
Compared with common wheat, ‘Certa’ appeared to be superior in
of Glu-D1 might be rearranged during the process of chromosomal
the measured photosynthetic characteristics at heading, flowering
recombination.
and early grain-filling stages (Table 2). Most of the lines derived from ‘Certa’ and JM47 showed improved SPAD value, Sc and Ci at all three growth stages. The Pn and Tr of these lines were a little lower than JM47 at heading stage, but increased rapidly at flowering stage and remained high at grain-filling stage when the photosynthetic capacity began to decrease in JM47, while the lines derived from ‘Certa’ and XN389 showed clearly higher SPAD value and Pn than XN389 at all three growth stages.
4 | DISCUSSION 4.1 | Chromosomal alterations and phenotypic variations in lines derived from hexaploid triticale and common wheat Hexaploid triticale carries the whole R chromosomes of rye, as well as the chromosomes of A genome and B genome of tetraploid
3.5 | Grain quality
wheat. In the process to transfer rye chromatin into common wheat using triticale as donor, recombination occurred between chromo-
The grain quality tested by near-infrared spectrometer showed that
somes from triticale and common wheat. In addition, rye chromo-
‘Certa’ had higher grain hardness and fibre content, but lower Zeleny
somes added to wheat background could induce modifications of
sedimentation and SDS sedimentation values than common wheat,
wheat chromosomes (Fu et al., 2013). In the present study, plentiful
13.75
12.00*
13.50
11.50*
12.50
12-4-2
12-5-1
12-6-2
12-10-1
12-11-4
22.00*
22.50*
21.00
22.00*
20.50
21.00
21.50*
20.00
20.00
28.50
20.50
17.00*
19.00*
18.50*
18.50*
19.00*
21.50
SN
71.00*
65.00
57.00
69.50*
55.50
52.50
55.00
52.00
53.00
102.00
65.00
67.50
62.50
54.00*
63.50
59.50
66.00
GNS
45.52*
48.30*
50.76*
47.33*
36.55*
45.96*
35.43*
48.28*
40.13
35.64
45.32*
41.29*
47.72
44.80*
48.06
44.74*
48.62
TGW (g)
77.10*
74.20*
74.80*
65.60
43.60*
60.00*
75.80*
54.80*
68.20
86.80
65.80
51.60*
51.00*
48.60*
41.20*
61.00*
75.20
BP (g)
31.05*
25.80*
27.00
27.20
18.00*
22.20*
22.40*
22.00*
30.00
33.00
23.00*
18.40*
20.40*
18.00*
17.40*
29.80
29.20
YP (g)
0.40*
0.35*
0.36*
0.41
0.41
0.37*
0.30*
0.40*
0.44
0.38
0.35*
0.36
0.40
0.37
0.42*
0.49*
0.39
HI
87.50*
82.00
95.75*
91.00*
76.50*
93.00*
91.50*
84.75*
80.25
139.75
118.50*
94.00*
91.75*
96.25*
98.50
78.00*
101.50
PH (cm)
9.75*
9.00*
14.25*
11.50
8.15*
8.10*
11.50
12.50
12.00
15.00
12.00
11.25*
11.00*
11.25*
12.75
11.75
13.00
SL (cm)
18.50*
16.38*
20.67
21.83
13.07*
13.88*
19.33*
20.83
19.83
20.55
23.38
23.88
21.63*
19.00*
19.20*
21.45*
23.38
FLL (cm)
1.90*
1.90*
2.20
2.13
1.80*
1.90
1.83*
1.83*
2.05
1.70
1.70
1.90*
1.78
1.52*
1.65
1.61*
1.78
FLW (cm)
24.75*
21.82*
31.89
32.66*
16.49*
18.43*
24.82*
26.85
28.47
24.49
28.11
31.74*
26.90*
20.55*
22.23*
24.35*
29.05
FLA (cm2)
195.00
196.00
196.00
199.00
196.00
196.00
198.00
188.00
199.00
194.00
198.00
200.00
192.00
192.00
192.00
192.00
197.00
DH (d)
202.00
203.00
203.00
205.00
202.00
202.00
205.00
198.00
206.00
204.00
203.00
205.00
199.00
201.00
199.00
199.00
204.00
DF (d)
2
2
1
1
1
1
1
1
0;
0;
2
2
2
2
2
2
4
IT
TN, tiller number; SN, spikelet number; GNS, grain number per spike; TGW, 1000-grain weight; BP, biomass per plant; YP, grain yield per plant; HI, harvest index; PH, plant height, SL, spike length; FLL, flag leaf length; FLW, flag leaf width; FLA, flag leaf area; DH, days to heading; DF, days to flowering; IT, infection type to stripe rust. *Significant at p = .05.
12.00*
12-4-1
14.25
Certa
16.00
20.00*
10-6-1
13.50
12.50*
10-3-1
12-2-1
16.50
7-5-1
12-1-2
14.25*
7-4-2
15.25
11.00*
7-3-1
XN389
12.00*
7-2-1
Certa/XN389
17.00
JM47
Certa/JM47//JM47
TN
Line
Cross
T A B L E 1 Agronomic traits of lines derived from Certa/JM47//JM47 and Certa/XN389
6
| LIU ET AL.
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ET AL.
F I G U R E 5 Agronomic traits of lines derived from Certa and common wheat. (a) Lines derived from Certa/JM47//JM47; (b) lines derived from Certa/XN389
|
7
51.85*
52.20*
52.30*
52.00*
50.15*
48.25
51.95*
12-4-1
12-4-2
12-5-1
12-6-2
12-10-1
12-11-4
53.35
Certa
53.85*
49.60
10-6-1
12-2-1
48.00
10-3-1
12-1-2
49.30
7-5-1
46.45
46.65
7-4-2
XN389
53.05*
7-3-1
45.60
54.85*
48.80
50.95*
51.90*
52.80*
51.00*
54.65*
57.30*
43.95
54.00
48.55*
49.20*
47.90
46.35
49.65*
47.45
49.55
58.55*
51.10
54.55*
53.70*
58.80*
55.75*
56.20*
57.25*
49.40
55.15
53.45*
49.75
52.05*
51.00
52.65*
51.05
19.18
18.93
22.73*
20.98*
20.87*
21.94*
20.62*
21.74*
20.22*
16.20
20.33
21.44
15.03*
18.48
16.45*
16.52*
19.43
17.90
19.21
20.38*
18.49
19.00
21.61*
23.69*
21.03*
18.05
24.14
19.92
22.61*
23.37*
24.86*
22.46*
21.65
19.61
Z65
15.74
19.74*
19.99*
18.28
22.47*
18.81*
18.06
19.70*
16.76
21.25
17.80
19.48*
21.84*
16.39
21.36*
19.43*
16.21
Z70
0.61
0.57
0.78*
0.84*
0.81*
0.71
0.67
0.64
0.62
0.81
0.52
0.81
0.90*
0.72
0.95*
0.80
0.65
Z55
0.36*
0.50
0.52
0.24*
0.49
0.26*
0.29*
0.50
0.58
0.76
0.44
0.48
0.68*
0.62*
0.61*
0.63*
0.49
Z65
Sc (mol m2 s1)
0.39*
0.31*
0.75
0.20*
0.78
0.58
0.20*
0.71
0.65
0.83
0.58
0.70*
0.68*
0.55
0.66*
0.65*
0.47
Z70
304.70*
283.31*
303.30*
314.20
308.61
309.59
303.36*
310.66
326.06
326.03
294.69
339.18*
335.05*
336.41*
343.19*
329.35
319.01
Z55
263.80
282.65
279.56
233.94*
282.68
209.57*
229.97*
278.99
293.28
293.61
271.61
263.75
286.84
278.36
286.03
291.35*
277.71
Z65
Ci (lmol m2 s1)
291.95
257.95*
306.12
223.42*
304.12
315.29
230.59*
313.50
317.38
318.24
313.49*
307.98
299.13
309.45*
300.84
309.45*
303.65
Z70
Pn, net photosynthetic rate; Sc, stomatal conductance; Tr, transpiration rate; Z55, heading stage; Z65, flowering stage; Z70, early grain-filling stage. *Significant at p = .05.
Certa/XN389
47.25
48.95
JM47
Certa/JM47//JM47
Z70
Z55
Z65
Z55
7-2-1
Line
Cross
Pn (lmol m2 s1)
SPAD value
T A B L E 2 Photosynthetic characteristics of lines derived from Certa/JM47//JM47 and Certa/XN389 at different growth stages
3.46
3.28
3.73
4.29*
4.18*
4.01*
3.65
3.76
3.24
4.10
3.00*
3.63
3.95
3.81
4.48
3.96
4.14
Z55
3.97
4.89
4.83
3.15*
4.77
3.16*
3.74
5.08*
4.66
5.94
4.41
4.47
5.60*
5.64*
5.58*
5.64*
4.45
Z65
Tr (mmol m2 s1)
4.00
3.60
5.22*
2.65*
5.50
4.01
2.75*
5.44*
4.38
5.46
4.94*
4.60
5.29*
4.70
5.27*
4.87*
4.04
Z70
8
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9
T A B L E 3 Grain quality traits and HMW-GS variations (Glu-1 loci) of lines derived from Certa/JM47//JM47 and Certa/XN389
Combination
Code
Protein (%)
Certa/ JM47//JM47
JM47
10.41
Fibre (%)
Gluten (%)
65.88
3.11
23.70
Glu-1 loci
Hardness
Zeleny (mL)
SDS (mL)
Glu-A1
Glu-B1
Glu-D1
Glu-R1
24.95
20.16
64.79
Null
7+9
3+12
–
7-2-1
9.95*
65.46
3.02*
21.89*
21.82*
22.23*
72.60*
Null
7+9
3+12
–
7-3-1
11.71*
64.34*
3.23*
27.79*
32.88*
24.43*
70.25*
Null
7+9
3+12
–
9.88*
66.11
3.09
21.96
23.54
21.59*
66.64
Null
7+9
3+12
–
7-5-1
10.46
66.08
3.11
23.78
22.65
22.55*
63.97
Null
7+9
3+12
–
10-3-1
10.31
64.56
3.11
22.93*
26.21
18.77*
60.96
Null
7+9
3+12
–
67.24*
3.09
22.31*
18.37*
17.90*
58.83*
Null
7+9
3+12
–
7-4-2
10-6-1
Certa/XN389
Starch (%)
9.99*
Certa
11.70
64.23
3.22
27.45
37.57
15.79
52.48
1
7+18
–
6+13
XN389
12.04
62.22
3.12
28.43
26.95
25.88
57.16
1
7+9
2+12
–
12-1-2
9.79*
66.08*
3.01*
20.89*
14.31*
20.07*
55.36
Null
–
2+12
6+13
12-2-1
11.12*
64.36*
3.26*
25.09*
18.28*
23.32*
60.78
1
7+9
2+12
–
12-4-1
10.01*
64.32*
3.21
21.68*
16.18*
20.63*
54.57
1
7+9
2+12
–
12-4-2
10.46*
63.85*
3.33*
23.45*
20.71*
20.56*
57.21
1
7+9
2+12
–
12-5-1
11.08*
64.84*
3.12
24.85*
18.37*
25.65
63.40*
1
7+9
2+12
–
12-6-2
10.34*
66.11*
3.05*
22.99*
18.29*
24.73
63.47*
1
7+9
4+12
–
12-10-1
10.73*
63.02*
3.34*
24.20*
21.16*
18.98*
51.44
1
7+9
2+12
–
12-11-4
9.62*
65.99*
3.12
20.68*
10.43*
23.77*
58.74
1
7+9
2+12
–
*Significant at p = .05. “–” indicates none.
recombination and structural alterations of wheat chromosomes
chromosomes were detected, supporting that 1R chromosome had
were identified. In addition to chromosome reorganization, great
higher transmission rate than other rye chromosomes.
morphological,
photosynthetic
and
HMW-GS
variations
were
detected in the progeny. For instance, novel patterns of HMW-GS that were different from those in the parents were observed in lines 12-1-2 and 12-6-2, indicating that the coding regions were rear-
4.3 | The potential of the new 1BL.1RS translocation lines
ranged, which remained to be studied further. Thus, in breeding pro-
The 1RS chromosome from S. cereale has been used in wheat
gramme involving distant hybridization, more attention should be
improvement over several decades. Although genes involving resis-
paid to the structural alterations of wheat chromosomes, which may
tance to stripe rust and powdery mildew located on the widely used
play an important role in wheat improvement (Li, Guo, Wang, & Ji,
1RS derived from Petkus rye are no longer completely effective to
2015; Li, Zhu, et al., 2015; Tang, Li, et al., 2014).
the new epidemic races, continuous efforts have been underway to search for new sources of 1RS that contain agronomically useful
4.2 | Occurrence frequency of rye chromosomes in progeny
genes. Several new 1BL.1RS translocation lines derived from different rye varieties were developed (Ko et al., 2002; Li, Guo, et al., 2015; Li, Zhu, et al., 2015; Tsuchida et al., 2008), and some were
In the present study, the occurrence frequency of rye chromosomes
secalin-absent (Gustafson et al., 2008; Masoudi-Nejad, Nasuda,
in the two populations was quite different even though both cross
McIntosh, & Endo, 2002). As several kinds of triticale and rye were
combinations using ‘Certa’ as the female parent, indicating that
involved in the breeding progress, the composition of rye chromo-
transmission rate of rye chromosomes was affected by the back-
somes of ‘Certa’ used in the present study could be complicated
ground of common wheat. In addition, the backcrossing in Certa/
(http://www.wheatpedigree.net/). In the present study, five 1BL.1RS
JM47//JM47 cross may decrease the occurrence frequency of rye
translocation lines derived from ‘Certa’ were identified, in which one
chromosomes. Ren, Lelley, and Robbelen (1991) also found that in
line (line 12-6-2) was found to lack the x-secalin locus. Compared
the F2 and F3 populations of octoploid triticale 9 common wheat,
with the wheat parent, grain starch content and SDS value were sig-
1R chromosome had the highest occurrence frequency while chro-
nificantly increased in line 12-6-2. Its potential in quality improve-
mosomes 6R and 7R showed the lowest frequency in progeny. Simi-
ment needs to be further studied. In practice, it was found that
lar phenomena were found in the F5 population derived from
1BL.1RS chromosome always paired as rod bivalents at diplotene
octoploid triticale 9 common wheat (Fu, Tang, & Ren, 2011) and
and diakinesis stages during meiosis, and thus, chiasmata and cross-
introgression lines of wheat 9 S. africanum Stapf (Lei, Li, Liu, & Yang,
ing over were difficult to form between 1RS and 1BS chromosomes,
2012). In Certa/XN389 cross in this study, only 1R and 1RS
so that the occurrence frequency of translocation without secalin
10
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chromatin was greatly decreased (Graybosch, Peterson, & Chung, 1999; Zhou et al., 2004). Secalin introduced by 1RS chromosome had negative effect on grain quality. In addition, the absence of 1BS aggravated that situation due to the loss of related genes encoding low molecular weight glutenin in 1BL.1RS translocation lines. Previous studies showed that some HMW-GS with high quality, for instance 1Dx5+1Dx10, could compensate for the negative effect caused by 1RS and improve grain quality in 1BL.1RS translocation lines (Wang, Ji, Wang, Zhang, & Zhang, 2006). The structural changes in wheat chromosomes in the present study represent a new genetic variation of wheat genome. Although little was improved in grain quality in the five 1BL.1RS translocation lines compared with their parents, there were wide variations in agronomic traits among which the photosynthetic capability was significantly increased. These 1BL.1RS translocation lines might have the potential to positively impact on wheat improvement by crossing with high-yielding germplasm or cultivars.
ACKNOWLEDGEMENTS This work was financially supported by the subproject of the 863 Program (2013AA102902) of the Ministry of Science and Technology, the China 111 Project (B12007) of the Ministry of Education of China; Science and Technology Innovation Team Plan of Shaanxi Province (2014KCT-25); and the Open Project of the State Key Laboratory of Crop Stress Biology for Arid Areas (CSBAA2017006).
CONFLICT OF INTEREST The authors declare that they have no conflict of interest. ORCID Caiyun Liu
http://orcid.org/0000-0001-7987-703X
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SUPPORTING INFORMATION Additional Supporting Information may be found online in the supporting information tab for this article.
How to cite this article: Liu C, Song Q, Zhang H, Yang Z, Hu Y-G. Molecular cytogenetic characterization and phenotypic evaluation of new wheat–rye lines derived from hexaploid triticale Certa 9 common wheat hybrids. Plant Breed. 2017;00:1–11. https://doi.org/10.1111/pbr.12523
