Scientia Agricultura Sinica ›› 2026, Vol. 59 ›› Issue (4): 723-733.doi: 10.3864/j.issn.0578-1752.2026.04.002

• CROP GENETICS & BREEDING·GERMPLASM RESOURCES·MOLECULAR GENETICS • Previous Articles     Next Articles

Development and Field Evaluation of Glyphosate-Resistant Wheat Germplasm Generated Through EMS Mutagenesis

CUI ShiYou1(), CHEN PengJun2, MIAO YuanQing2, HAN JiJun2, SHEN JunMing1   

  1. 1 Nantong Ruide Agi-Tech Co. Ltd, Nantong 226008, Jiangsu
    2 Marine Geological Survey of Jiangsu Province, Nanjing 210007
  • Received:2025-10-15 Online:2026-02-10 Published:2026-02-10

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Abstract:

【Objective】This study aimed to develop novel glyphosate-resistant wheat germplasm using EMS mutagenesis to mitigate weed infestation in wheat fields. Resistant mutant plants were selected through field screening, and the mutation profiles of the 5-enolpyruvylshikimate-3-phosphate synthase (EPSPS) gene as well as optimal application conditions were characterized, offering a practical approach for breeding glyphosate-resistant wheat varieties.【Method】A mutant population was generated by treating newly germinated seeds of Zhenmai 9 with EMS mutagenesis. Resistant mutants were isolated through multiple rounds of glyphosate screening in the field across M2 and M3 generations. Promising lines, including GR1, GR19, and GR23, were identified via pedigree selection, combined with yield and resistance phenotype screening. Mutation sites in the EPSPS gene were detected by PCR amplification and sequencing, while expression levels of TaEPSPS-4A, TaEPSPS-7A, and TaEPSPS-7D were analyzed using RT-qPCR. Field trials involving different glyphosate doses and application growth stages were conducted to systematically evaluate herbicide efficacy and determine the appropriate dosage and timing for safe application.【Result】The resistance mutant frequency in the M2 population was 6.53×10-6. In the M3 generation, 43 mutant plants exhibiting tolerance to 4× the recommended glyphosate dose were successfully obtained. Sequencing analysis revealed that resistant lines GR1 and GR19 harbored 5 and 3 mutation sites in TaEPSPS-7D, respectively, whereas GR23 carried 5 mutation sites in TaEPSPS-4A. Expression analysis indicated that glyphosate treatment significantly downregulated most homoeolog genes in the three mutation lines, irrespective of whether those genes carried resistance mutations. Field trials demonstrated 100% weed control efficacy across all glyphosate treatments, significantly superior to isoproturon. As the glyphosate doses increased, wheat seedling height and fresh weight showed a decreasing trend, but most differences with the untreated control were not significant, indicating no substantial adverse effects on growth. Yield analysis revealed that treatment with 1× and 2× doses did not cause significant yield reduction, whereas 4× and 8× doses led to significant reductions of 3.04% and 4.63%, respectively. Growth stage-specific trials further indicated that spraying a 2× dose of glyphosate from seedling to jointing stages had no significant impact on plant growth, but application at the booting stage significantly reduced plant height, fresh weight, and grain yield, resulting in a 6.48% yield loss.【Conclusion】The combination of EMS mutagenesis and field screening successfully generated new glyphosate-resistant wheat germplasm capable of withstanding 4× the recommended glyphosate dose. Multiple point mutations in the non-active center of the EPSPS enzyme confered enhanced glyphosate resistance without compromising yield. For practical application of such resistant varieties, the optimal weed control window is during wheat green-up (early March), using 41% glyphosate isopropylamine salt at 840-1 680 g ae·hm-2, diluted in 450 L·hm-2 of water, applied as foliar spray to weeds under rain-free conditions.

Key words: glyphosate resistance, wheat (Triticum aestivum), EMS mutagenesis, mutation site, gene expression, field efficacy trial

Table 1

Primers used for homologous PCR amplification of wheat EPSPS gene"

引物Primer 序列Sequences (5′-3′)
共同正向引物Common homoeolog EPSPS forward primer ACAGTGAGGATGTCCACTACATGCTTGA
7A反向引物Chr.7A EPSPS homoeolog reverse primer ACTTCTCTGACAGAGAACAGAAGTGTGCAC
4A反向引物Chr.4A EPSPS homoeolog reverse primer TTGTGTAAGGTCGCATTGATCGTACTACCA
7D反向引物Chr.7D EPSPS homoeolog reverse primer GAAAACTAGAATCATGCTTTTGTACTCCACTATC

Table 2

Genome-specific primers performed in RT-qPCR analysis of the EPSPS transcripts"

基因Gene 正向引物Forward primer (5′-3′) 反向引物Reverse primer (5′-3′)
TaEPSPS-4A TACTTGAGATGATGGGAGCG GCAACAACGGCAAGAGTCATT
TaEPSPS-7A GAACATCACGGCGATCGAC TCTTTCGGGTGCATCCAGGG
TaEPSPS-7D GAACGTCACGGCGATCGAT TCTTTCTGGTGCACCCCGGA
TaActin GTTGGTGATGAGGCCCAATC GTGCTACACGGAGCTCATTG

Fig. 1

Field screening tests of glyphosate-resistant mutants A: Field plots of wild type (WT) Zhenmai 9; B: GR mutant plants of M3 generation obtained spraying 4×glyphosate; C: Field plots of GR mutant line GR23 (M3:5) with 8× glyphosate treatment"

Table 3

Yield performance of 7 glyphosate resistant mutant lines (M3:4) and its resistance to glyphosate"

突变系
Mutant line		 籽粒产量Grain yield (kg·m-2) 对草甘膦的抗性
Resistance to glyphosate
喷草甘膦 With glyphosate 不喷草甘膦 Without glyphosate P值 P value
GR1 0.765±0.055 0.695±0.120 0.116 抗Resistant
GR2 0.525±0.019 0.648±0.017 0.000 抗Resistant
GR3 0.655±0.101 0.788±0.073 0.042 抗Resistant
GR4 0.548±0.032 0.585±0.063 0.175 抗Resistant
GR5 0.672±0.139 0.712±0.116 0.336 抗Resistant
GR6 0.462±0.079 0.407±0.021 0.130 抗Resistant
GR9 0.713±0.024 0.755±0.028 0.034 抗Resistant
GR19 0.779±0.017 0.748±0.020 0.144 抗Resistant
GR23 0.636±0.015 0.625±0.019 0.686 抗Resistant

Fig. 2

Ananlysis of mutation sites in glypphosate-resistant mutant line GR1, GR19 and GR23 The sequences marked in gray are exons, and the unmarked sequences are introns; the sequences marked in purple correspond to the base sequences of the EPSPS enzyme active center; the mutated codons are marked in bold red, and the underlined bases indicate the mutation sites; and the blue characters indicate the amino acid substitutions caused by mutations"

Fig. 3

Expression analysis of three wheat EPSPS homoeologous genes of the mutant lines GR1, GR19 and GR23 Different letters of the same gene indicate significant differences between treatments (P<0.05)"

Table 4

The effect of herbicide treatment on wheat shoot height, shoot fresh weight, and weed control"

药剂
Herbicide		 剂量
Dose
(g ae·hm-2)		 GR1 GR19 GR23
苗高
Shoot height
(cm)		 苗鲜重
Shoot fresh weight
(cm)		 株防效
Control effect of plant (%)		 苗高
Shoot height
(cm)		 苗鲜重
Shoot fresh weight
(cm)		 株防效
Control effect of plant (%)		 苗高
Shoot height
(cm)		 苗鲜重
Shoot fresh weight
(cm)		 株防效
Control effect of plant (%)
草甘膦1)
Glyphosate		 840 42.3a 2.98a 100.0a 38.9b 2.88a 100.0a 42.0a 2.95a 100.0a
1680 43.0a 2.73b 100.0a 41.3ab 2.77b 100.0a 39.3b 2.72b 100.0a
3360 40.7ab 2.72b 100.0a 44.5a 3.00a 100.0a 41.2ab 2.65b 100.0a
6720 41.3a 2.69b 100.0a 40.6b 2.82a 100.0a 38.9b 2.74b 100.0a
异丙隆2) Isoproturon 2250 40.7ab 2.67b 77.0b 38.9b 2.71b 73.7b 40.0ab 2.71b 76.7b
空白对照3) CK - 40.5ab 2.71b - 41.1ab 2.99a - 40.9ab 2.78b -

Table 5

The effect of herbicide treatment on yield of wheat mutant lines GR1, GR19 and GR23"

药剂 Herbicide 剂量 Dose (g ae·hm-2) GR1 GR19 GR23 平均 Average
草甘膦Glyphosate 840 0.743a 0.777a 0.741a 0.754a
1680 0.730ab 0.759ab 0.734ab 0.741b
3360 0.721bc 0.749b 0.729bc 0.733c
6720 0.713c 0.739b 0.712c 0.721d
异丙隆Isoproturon 2250 0.717bc 0.745b 0.717bc 0.726cd
空白对照CK - 0.745a 0.779a 0.744a 0.756a
平均Average 0.728b 0.758a 0.730b

Table 6

The effect of glyphosate treatment at different growth stages on the growth of seedlings and yield of wheat mutant line GR1"

生育期
Growth stage		 处理
Treatment		 产量及构成因素Yield and its components
苗高
Shoot height
(cm)		 单株鲜重
Fresh weight per plant (g)		 单株分蘖数
Tillering No. per plant		 籽粒产量
Grain yield
(kg·m-2)		 穗数
Spike No.
(m-2)		 每穗粒数
Grain No. per spike		 千粒重
1000-grain weight (g)
苗期
Seedling stage		 草甘膦Glyphosate 11.7a 0.49a 2.03a 0.724a 380.7a 41.1a 46.1a
CK 13.2a 0.42a 2.12a 0.725a 379.6a 41.6a 45.7a
分蘖期
Tillering stage		 草甘膦Glyphosate 21.3a 2.35a 3.15a 0.721a 378.9a 41.5a 45.8a
CK 21.5a 2.37a 3.11a 0.726a 381.6a 41.3a 45.7a
拔节期
Jointing stage		 草甘膦Glyphosate 60.2a 10.5a 3.23a 0.723a 379.5a 41.5a 45.6a
CK 61.4a 10.6a 3.58a 0.732a 383.2a 42.3a 45.3a
孕穗期
Booting stage		 草甘膦Glyphosate 67.1b 11.5b - 0.678b 376.6a 38.8b 45.1a
CK 71.3a 12.7a - 0.725a 378.1a 41.7a 45.9a
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