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100 Gy [sup.60]Co [gamma]-ray induced novel mutations in tetraploid wheat.

1. Introduction

Human activities and natural calamities decreased the biological diversity and narrowed the genetic variability that limits crop breeding. Novel mutations in plants, which are crucial for improving resistance/tolerance to environmental stress, enhancing quality and yield traits, and facilitating the seed set of hybrid, have been created, such as in Arabidopsis [1], rice [2], maize [3], wheat [4], and some horticultural plants [5].

Since the 1970s, [gamma]-rays, sodium azide, and ethyl methane sulfonate (EMS) have been used for wheat breeding [4]. Inducing mutation with [sup.60]Co [gamma]-ray is an effective way and had bred some hexaploid wheat cultivars. Guinness/1322 (Bulgaria), for an example, was mutationally bred from Katya (a hexaploid wheat cultivar from Bulgaria) by 50 Gy [sup.60]Co [gamma]-ray [6]. Compared with Katya, Guinness/1322 shows better lodging and shedding resistance, better ecological adaptability of drought tolerance, and higher productivity [7]. Inducing mutation with [sup.60]Co [gamma]-ray was also used for tetraploid wheat breeding, but only two cultivars, Yavor (Bulgaria) and Implus (Turkey), were bred from durum wheat (AABB, 2n = 4x = 28) and different frequencies of induced mutations were observed under 100 Gy [sup.60]Co [gamma]-ray [7, 8].

Tetraploid wheat (AABB, 2n = 4x = 28) distributes widely and adapts extensively to the environment and contains considerable wealth of genetic and morphological variation [9], such as high abilities of powdery mildew resistance in Triticum dicoccoides Korne [10], abundant genetic diversity of storage proteins in T dicoccoides [10] and Triticum turgidum L. [11], valuable genes contributing to the grains per spike in Triticum carthlicum Nevski [12], dwarf genes in Triticum polonicum L. [13], and high content of gluten and tolerance to the saline in Triticum durum Desf. [14, 15]. Tetraploid wheat with AB genomes is important natural resources for breeding [16]. Therefore, creating novel mutation through radiation in tetraploid or hexaploid wheat may be an effective way for wheat breeding.

In the present study, 10 accessions of tetraploid wheat were radiated with 100 Gy [sup.60]Co [gamma]-ray. Following the radiation, mutations of the agronomic traits, cytogenetics and high-molecular-weight glutenin subunits (HMW-GSs) were observed, which could be used for further selection and utilization of the radiated progenies.

2. Materials and Methods

2.1. Materials. All seeds of the accessions were deposited at Triticeae Research Institute, Sichuan Agricultural University, Sichuan, China. Information of the accessions was listed in Table 1.

2.2. Radiation. 20 seeds of each accession were radiated with 100 Gy [sup.60]Co [gamma]-ray at the Institute of Biological and Nuclear Technology, Sichuan Academy of Agricultural Sciences, China. Dose rate was 1.1Gy/min, and unirradiated seeds were used as a control (CK).

2.3. Seed Germination. Respective 20 radiated and CK seeds of each accession were exposed with 4[degrees]C for 24 hours and germinated with 25[degrees]C. The germination energy (percentage of the seeds germinated in 10 days) and germination rate were calculated as follows:

Germination rate = Number of germinated seeds/Total number of seeds (20) x 100%. (1)

2.4. Agronomie Characters Identification. Radiated seedlings and CK ones were planted in the field. Agronomic characters including plant height, tiller number, seed set, and other special characters (secondary tillering, stalk with wax powder, and dwarf) were observed.

2.5. Meiotie Analysis. Young spikes were fixed in Carnoy's solution II (ethanol:chloroform:acetic acid = 6:3:1V/V) and stored at 4[degrees]C. The pollen mother cells were stained with improved phenol fuchsin. Observations of the chromosome pairing of meiosis were made and documented with an Olympus BX-51 microscope coupled with a Photometric SenSys CCD camera. 60 cells of each accession were counted to confirm the pairing in the meiotic process.

2.6. High-Moleeular-Weight Glutenin Subunits (HMW-GSs). HMW-GSsofradiatedand CK seedswereanalyzedaccording to the method of Wan et al. [17]; eight seeds were tested in every single plant. The HMW-GSs of Chinese Spring (Null, 7 + 8, 2 + 12) were used as marker.

3. Results

The results of seed germination were shown in Table 2. Among all the 10 accessions, Tritieum dieoeeum Schrank (PI434999) with only 1 seed germinated (5%) is the most sensitive to this dose of radiation. T. polonieum (As304) and T. earthlieum (As293) are 35% and 40% of germinations, respectively. The germinated ratio of the other accessions varied from 80% to 95%.

The agronomic characters of all the radiated accessions were observed in the field. The results of varied agronomic characters of 4 radiated plants were shown in Table 3 and Figure 1. Compared with CK, T. dicoccoides (As838) 2397 had 11 secondary tillerings and stalk with wax powder (Figure 1(a)). Dwarfs were observed in both T. turgidum (As2255) 253-10 (Figure 1(b)) and T. polonieum (As302) 224-14 (Figure 1(d)). Average height of T. turgidum (As2255) CK is 103.2 [+ or -] 2.5 cm, while that of T turgidum (As2255) 253-10 is 68.5 cm. Average height of T. polonieum (As302) CK is 145.7 [+ or -] 5.9 cm, while that of T. polonieum (As302) 224-14 is 98.1 cm. T. earthlieum (As293) 250-1 was senescence, withered before harvest (Figure 1(c)). No other mutational agronomic characters were observed in other plants.

Chromosome pairing in meiosis of the plants with varied agronomic characters and some other radiated plants were also observed (Table 4). Univalents, trivalents, quadrivalents, and lagging chromosomes in meiosis were detected in few cells of the observed accessions. Quadrivalent was observed in T. dicoccoides (As835) 237-9 (Figure 2(a)). Univalents were observed in T. dicoccoides (As835) 237-11 (Figure 2(b)) and in T dicoccoides (As838) 239-7 (Figure 2(c)). Trivalent in T. dicoccoides (As838) 239-8 was observed (Figure 2(d)). The normal chromosome pairing was shown in Figure 2(e). Lagging chromosomes were observed in T. polonieum (As304) (Figure 2(f)). Chromosome pairing results were as follows: radiation treatment had no effect on meiosis of 3 individuals with varied agronomic characters, T. earthlieum (As293) 250-1, T polonieum (As302) 224-14 and T. turgidum (As2255) 253-10. Their chromosome pairing were 2n = 28 = 13.27II (ring) + 0.73II (rod), 2n = 28 = 10.56II (ring) + 3.43II (rod) and 2n = 28 = 13.05II (ring) + 0.95II (rod), respectively. Meanwhile, chromosome pairing of T dicoccoides (As838) 239-7 with 2n = 28 = 0.44I + 12.26II (ring) + 1.32II (rod) + 0.08III, exhibited a trait of 11 secondary tillerings and stalk with wax powder. The interference of chromosome pairing were also observed in radiated plants T. dicoccoides (As835) 237-11 (2n = 28 = 0.39I + 12.44II(ring) + 1.36n(rod))and T. dicoccoides (As835) 237-9 (2n = 28 = 0.44I + 11.79II (ring) + 1.05II (rod) + 0.2im + 0.53IV) with 1Ax silence in seed numbered T. dicoccoides (As835) 237-9-5. T. dicoccoides (As838) 239-8 (2n = 28 = 1I + 12.32II (ring) + 0.64II (rod) + 0.36III) with 1Ax silence in seed numbered T dicoccoides (As838) 239-8-2 and T polonieum (As304) 230-7 2n = 28 = 0.10I + 12.11II (ring) + 1.41II (rod) + 0.10III + 0.18IV with a novel HMW-GS observed in seed numbered T polonieum (As304) 230-7-1. Their chromosome pairings of meiotic process are abnormal, compared to the meiosis of the CK ones.

HMW-GSs of eight randomly selected seeds of each single radiated plant of all the 10 tetraploid accessions were tested by SDS-PAGE. Three mutations were found. 1Ax was silent in T dicoccoides (As835) 237-9-5 and T dicoccoides (As838) 239-8-2 (Figures 3(a) and 3(b)). Compared with CK, a novel HMW-GS in T. polonieum (As304) 230-7-1 was detected whose electrophoretic mobility was between 1By8 and 1Dy12 which were the HMW-GSs of Chinese Spring (Figure 3(c)).

4. Discussion

Different species has different suitable dose of radiation intensity, such as 300 to 700 Gy [sup.60]Co [gamma]-ray in Sorghum bieolor (L.) Moench [18], less than 200 Gy in Roegneria [19]. Suitable dose of radiation is various among different species in same genus [19]. In the present study, 100 Gy [sup.60]Co [gamma]-ray differently induced mutations in the tetraploid wheat accessions. The results of germination energy and germination rate suggest that T. dicoccum (PI434999) is the most sensitive to the treatment and this dose of radiation is lethal dose to it. As to most tetraploid wheat, 100 Gy [sup.60]Co [gamma]-ray radiation is an insufficient dose to induce mutation.

Chromosomal translocation, chromosome breakage, and deletions in chromosome which came from radiation mutation may lead to defects in chromosome pairing [20]. In this study, the abnormal chromosome pairings in meiotic process, such as univalents, trivalents, quadrivalents, and lagging chromosomes, were observed in pollen mother cells of some radiated plants, suggesting that 100 Gy [sup.60]Co [gamma]-ray might create some mutations at chromosome level. Univalents were observed in T dicoccoides (As838) 239-7 with 11 secondary tillerings and stalk with wax powder. Meanwhile, quadrivalent in T dicoccoides (As835) 237-9, trivalents in T. dicoccoides (As838) 239-8, and lagging chromosomes in T polonicum (As304) 230-7 were observed with HMW-GS mutations. Thus, the abnormal chromosome pairing of meiotic process reflected radiation mutations. Meiotic process observation could be used as a tool for mutation identification at wheat earing stage.

HMW-GSs are important storage proteins in wheat and its related species and 10% of endosperm proteins are HMWGSs [21-23]. Theoretically, tetraploid wheat should contain 4 different HMW-GSs, 1Ax, 1Ay, 1Bx and 1By [23], but only one or two, no more than three subunits, were expressed due to gene silencing. Different HMW-GSs combinations have different effect on flour quality [23]. In the present study, compared with CK, 1Ax was silent in T dicoccoides (As835) 237-9-5 and T. dicoccoides (As838) 239-8-2. HMW-GS gene silencing might be caused by specific nucleotide substitutions in the promoter region [21] and single repeat changes or repeat indels or large deletions in codon region [22, 23]. A novel HMW-GS was detected in T. polonicum (As304) 230-7-1 and its electrophoretic mobility was between 1By8 and 1Dy12 which were the HMW-GSs of Chinese Spring. Single nucleotide mutation or repeat deletions could restore the expression of genes; homoeologous recombination might be a novel pathway for allelic variation or molecular evolution of HMW-GSs [22, 24]. The mechanism of mutations in HMWGS is under research.

Dwarf genes were found to be affecting architecture of rice plant [25]. GID1 gibberellin receptors affect the plant height of Arabidopsis [26]. 10 dwarfing genes/alleles have been discovered from tetraploid wheat [13, 27-29]. Associating with an extreme dwarf trait, only a few dwarfing genes have been used for wheat breeding worldwide [30]. Digging new plant height reducinggene is more and more important for wheat dwarf breeding. In the present study, significant plant dwarf was observed in both radiated plants T. polonicum (As302) 224-14 and T turgidum (As2255) 253-10. The average height of T. polonicum (As302) is 145.7 [+ or -] 5.9 cm, while the radiation mutation of T. polonicum (As302) 224-14 is 98.1 cm in height. A nature mutant dwarf accession of T. polonicum (As304) is 68 cm in height. Radiation may cause different dwarf gene and the effect of dwarf accumulated T turgidum 253-10 shows an extreme dwarf trait.

Inducing mutations for genetic improvement in breeding resources has been successfully and widely used for plant breeding. Sodium azide, EMS, and [gamma]-rays are major tools for mutation. Sodium azide was widely used for mutation and breeding in rice [31], barley [32], tomato [33], and maize [3] but was not an effective mutagen in Arabidopsis [34]. EMS mainly induced single nucleotide mutations in Arabidopsis thaliana [1], hexaploid wheat, and Triticale [35]. Inducing mutations through chromosome aberration and single nucleotide mutant enriched the gene banks of the species [33]. During the past fifty years, about 130 wheat cultivars bred from mutation have been widely produced in China [36]. In the present study, some novel mutations in several tetraploid wheat cultivars were induced by 100 Gy [sup.60]Co [gamma]-ray, such as HMW-GS and dwarf trait, which could be used as resources for theoretical study and future wheat breeding.

5. Conclusion

In the present study, 100 Gy [sup.60]Co [gamma]-ray differently induced mutations in accessions of tetraploid wheat. Following the radiation the germinated ratio of the materials varied from 5% to 95% and this dose of radiation is lethal dose to T. dicoccum (PI434999). The effects of radiation on the meiotic process of pollen mother cells and HMW-GSs were observed. Univalents, trivalents, quadrivalents, and lagging chromosomes in meiosis were detected in few cells of the observed accessions. As to HMW-GS, 1Ax was silent in T. dicoccoides (As835) 237-9-5 and T. dicoccoides (As838) 2398-2 and a novel HMW-GS was detected in T. polonicum (As304) 230-7-1 whose electrophoretic mobility was between 1By8 and 1Dy12 which were the HMW-GSs of Chinese Spring. Compared to the CK, T. dicoccoides (As838) 239-7 had 11 secondary tillerings and stalk with wax powder. Plant dwarfs were also observed; the height of the radiated T. turgidum (As2255) 253-10 is 68.5 cm and T. polonicum (As302) 224-14 is 98.1 cm. These mutations would be resources for the future wheat breeding.

http://dx.doi.org/ 10.1155/2014/725813

Conflict of Interests

The authors declare that there is no conflict of interests regarding the publication of this paper.

Acknowledgments

This work was supported by the National Natural Science Foundation of China (nos. 31301349,30870154,30901052, and 30900087), the Bureau of Education of Sichuan Province (13ZA0256), and the Bureau of Science and Technology of Sichuan Province, China.

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Chuntao Yang, (1,2) Jianshu Zhu, (1) Yun Jiang, (3) Xiaolu Wang, (1) Mengxue Gu, (1) Yi Wang, (1) Houyang Kang, (1) Xing Fan, (1) Lina Sha, (1) Haiqin Zhang, (1) Pu Xuan, (3) and Yonghong Zhou (1,2)

(1) Triticeae Research Institute, Sichuan Agricultural University, Wenjiang, Chengdu, Sichuan 611130, China

(2) Key Laboratory of Crop Genetic Resources and Improvement, Ministry of Education, Sichuan Agricultural University, Wenjiang, Chengdu, Sichuan 611130, China

(3) Institute of Biological and Nuclear Technology, Sichuan Academy of Agricultural Sciences, Chengdu, Sichuan 610061, China

Correspondence should be addressed to Yonghong Zhou; zhouyh@sicau.edu.cn

Received 27 February 2014; Revised 25 April 2014; Accepted 27 April 2014; Published 20 May 2014

Academic Editor: Wujun Ma

TABLE 1: Materials used in this study.

Species                      Accession      Ploidy       Genome
                              number

Triticum carthlicum Nevski     As293     2n = 4x = 28     AABB
Triticum dicoccoides Korne     As835     2n = 4x = 28     AABB
Triticum dicoccoides Korne     As838     2n = 4x = 28     AABB
Triticum dicoccum Schrank    PI434999    2n = 4x = 28     AABB
Triticum durum Desf.           As781     2n = 4x = 28     AABB
Triticum polonicum L.          As302     2n = 4x = 28     AABB
Triticum polonicum L.          As304     2n = 4x = 28     AABB
Triticum turanicum Jakubz.    As2279     2n = 4x = 28     AABB
Triticum turgidum L.          As2255     2n = 4x = 28     AABB
Triticum turgidum L.           As313     2n = 4x = 28     AABB

Species                              Origin

Triticum carthlicum Nevski           Japan
Triticum dicoccoides Korne           Israel
Triticum dicoccoides Korne           Israel
Triticum dicoccum Schrank    Bosnia and Herzegovina
Triticum durum Desf.                America
Triticum polonicum L.           Xinjiang, China
Triticum polonicum L.           Xinjiang, China
Triticum turanicum Jakubz.      Xinjiang, China
Triticum turgidum L.             Beijing, China
Triticum turgidum L.            Jianyang, China

TABLE 2: The results of the germination.

Species                  Total number   Germination
                           of seeds       number

T. carthlicum (As293)         20             8
CK                            20            19
T. dicoccoides (As835)        20            18
CK                            20            20
T dicoccoides (As838)         20            18
CK                            20            20
T dicoccum (PI434999)         20             1
CK                            20            19
T. durum (As781)              20            17
CK                            20            20
T polonicum (As302)           20            16
CK                            20            20
T polonicum (As304)           20             7
CK                            20            20
T turanicum (As2279)          20            16
CK                            20            20
T turgidum (As2255)           20            17
CK                            20            20
T turgidum (As313)            20            19
CK                            20            20

Species                  Germination   Germination
                          rate (%)     energy (%)

T. carthlicum (As293)        40            35
CK                           95            95
T. dicoccoides (As835)       90            85
CK                           100           100
T dicoccoides (As838)        90            90
CK                           100           95
T dicoccum (PI434999)         5             5
CK                           95            95
T. durum (As781)             85            80
CK                           100           100
T polonicum (As302)          80            75
CK                           100           100
T polonicum (As304)          35            35
CK                           100           100
T turanicum (As2279)         80            80
CK                           100           100
T turgidum (As2255)          85            85
CK                           100           100
T turgidum (As313)           95            95
CK                           100           100

TABLE 3: Special agronomic characters of the materials under
radiation treatment.

Species                  Individuals   Plant height (cm)

T carthlicum (As293)        250-1             85.4
                             CK        105.4 [+ or -] 1.8
T. dicoccoides (As838)      239-7            124.7

                             CK        113.7 [+ or -] 3.7
T. polonicum (As302)       224-14             98.1
                             CK        145.7 [+ or -] 5.9
T turgidum (As2255)        253-10             68.5
                             CK        103.2 [+ or -] 2.5

Species                  Individuals   Tiller number   Seed set

T carthlicum (As293)        250-1           11           0.01
                             CK        8 [+ or -] 2      0.85
T. dicoccoides (As838)      239-7           16           0.79

                             CK        17 [+ or -] 1     0.51
T. polonicum (As302)       224-14            4           0.39
                             CK        3 [+ or -] 0      0.8
T turgidum (As2255)        253-10            7           0.53
                             CK        5 [+ or -] 2      1.67

Species                  Individuals      Special characters

T carthlicum (As293)        250-1          Plant senescence
                             CK
T. dicoccoides (As838)      239-7      11 secondary tillerings,
                                        stalk with wax powder
                             CK
T. polonicum (As302)       224-14               Dwarf
                             CK
T turgidum (As2255)        253-10               Dwarf
                             CK

TABLE 4: Chromosome pairing at metaphase I in the pollen mother cells
of the materials with special traits after radiation treatment.

Species       Individuals   Number of    Number of
                              cells     chromosomes
                            observed

T                250-1         60           28
carthlicum
(As293)
                  CK           60           28

T polonicum     224-14         60           28
(As302)

                  CK           60           28

T turgidum      253-10         60           28
(As2255)

                  CK           60           28

                 237-9         60           28
T
dicoccoides     237-11         60           28
(As835)

                  CK           60           28

                 239-7         60           28
T
dicoccoides      239-8         60           28
(As838)

                  CK           60           28

T polonicum      230-7         60           28
(As304)

                  CK           60           28

                                          Chromosomes pairing

Species       Individuals    I              II            III     IV

                                   Total   Ring    Rod

T                250-1                     13.27   0.73
carthlicum                         14.00
(As293)                                    11-14   0-3
                  CK                       13.55   0.45
                                   14.00
                                           11-14   0-3
T polonicum     224-14                     10.56   3.43
(As302)                            13.99
                                           10-13   1-4
                  CK                       11.08   2.92
                                   14.00
                                           10-13   1-4
T turgidum      253-10                     13.05   0.95
(As2255)                           14.00
                                           10-14   0-4
                  CK                       13.26   0.74
                                   14.00
                                           10-14   0-4
                 237-9      0.44           11.79   1.05   0.21   0.53
T                                  12.84
dicoccoides     237-11      0-2            9-13    0-3    0-1    0-1
(As835)                     0.39   13.80   12.44   1.36
                            0-4            10-14   0-3
                  CK                       13.15   0.85
                                   14.00
                                           11-14   0-3
                 239-7      0.44           12.26   1.32   0.08
T                                  13.58
dicoccoides      239-8      0-2            10-13   0-4    0-1
(As838)                      1     12.96   12.32   0.64   0.36
                            0-4            10-14   0-2    0-1
                  CK                       13.36   0.64
                                   14.00
                                           12-14   0-2
T polonicum      230-7      0.10           12.11   1.41   0.10   0.18
(As304)                            13.52
                            0-1            10-14   0-4    0-1    0-1
                  CK                       13.05   0.95
                                   14.00
                                           10-14   0-4
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Title Annotation:Research Article
Author:Yang, Chuntao; Zhu, Jianshu; Jiang, Yun; Wang, Xiaolu; Gu, Mengxue; Wang, Yi; Kang, Houyang; Fan, Xi
Publication:The Scientific World Journal
Article Type:Report
Date:Jan 1, 2014
Words:4337
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