Scientia Agricultura Sinica ›› 2026, Vol. 59 ›› Issue (12): 2551-2562.doi: 10.3864/j.issn.0578-1752.2026.12.002

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

Genetic Dissection of Stem Internode Length and Its Effects in Wheat Based on a Genome-Wide Association Study

YE MeiJin1,7(), WANG TongZhu2, CHEN Bin2, LOHANI Md Nahibuzzaman2, CHEN JiaTing2, HU XinRong2, YIN Li2, WANG Chao2, ZHANG HaoPeng2, YANG Xia2, WANG JiaLin3, YAO QiFu4, DING PuYang5, WANG Feng1, LI XiaoYu6(), MA Jian2()   

  1. 1 College of Chemistry and Life Sciences, Chengdu Normal University, Chengdu 611130
    2 Triticeae Research Institute, Sichuan Agricultural University, Chengdu 611130
    3 College of Agronomy, Sichuan Agricultural University, Chengdu 611130
    4 College of Agroforestry Engineering and Planning, Tongren University/Guizhou Key Laboratory of Biodiversity Conservation and Utilization in the Fanjing Mountain Region, Tongren 554300, Guizhou
    5 School of Biological and Pharmaceutical Sciences, Mianyang Teachers’ College, Mianyang 621000, Sichuan
    6 Nanchong Academy of Agricultural Sciences, Nanchong 637000, Sichuan
    7 Sichuan Provincial Key Laboratory for Development and Utilization of Characteristic Horticultural Biological Resources, Chengdu 611130
  • Received:2025-12-09 Accepted:2026-01-20 Online:2026-06-16 Published:2026-06-16
  • Contact: LI XiaoYu, MA Jian

Abstract:

【Objective】Plant height is a key agronomic trait in wheat that influences both yield potential and lodging resistance. It is primarily determined by the elongation of stem internodes. This study aimed to systematically evaluate the internode lengths of 224 Sichuan wheat cultivars, identify stable quantitative trait loci (QTL) regulating internode length and clarify their effects on agronomic traits, and screen the underlying candidate genes, thus providing important genetic resources and a theoretical basis for the targeted breeding of high-yield wheat varieties. 【Method】The lengths of the first (IL1), the second (IL2), and the third (IL3) internodes were measured under two different environments. The panel was genotyped using the wheat 120K SNP array. A genome-wide association study (GWAS) was employed to identify QTL regulating internode length. The phenotyping data was used for correlation analysis and the interpretation of the genetic effects of major QTL. The candidate genes of the major QTL were predicted based on wheat omics data. 【Result】Phenotypic analysis revealed continuous variation for all internode traits, with high broad-sense heritability estimates ranging from 75% to 89%. Correlation analysis showed that lengths of all three internodes were positively correlated with final plant height, with IL1 exhibiting the strongest correlation. Furthermore, IL2 and IL3 were highly correlated with each other, suggesting coordinated genetic regulation of the lower stem internodes. GWAS detected four stable QTLs on chromosomes 5A and 4D, namely QIL1.sau.5A for IL1, two tightly linked but distinct QTL QIL2.sau.5A.1 and QIL2.sau.5A.2 for IL2, and QIL3.sau.4D for IL3, respectively. Pleiotropy analysis demonstrated that QIL2.sau.5A.1 significantly increased plant height and spike length, while QIL3.sau.4D primarily promoted internode elongation and overall plant height. Based on functional annotation and spatiotemporal expression data, five candidate genes potentially involved in transcriptional regulation, hormone signal transduction, and cell growth were identified. 【Conclusion】This study elucidated the genetic architecture of internode elongation in a panel of Sichuan wheat cultivars and identified two novel QTLs and several pleiotropic loci. These stable QTLs and the underlying candidate genes provided valuable resources for molecular marker-assisted selection aiming at optimizing plant height, improving lodging resistance, and enhancing yield potential in wheat.

Key words: Triticum aestivum L., internode lengths, SNP markers, GWAS, agronomic traits, candidate genes

Table 1

Phenotypic assessment of the natural population"

性状
Trait
环境
Environment
最大值
Max
最小值
Min
平均值
Mean
标准差
Standard deviation
变异系数
CV (%)
广义遗传力
H2 (%)
第一节间长度
IL1 (cm)
WJ2025 49.16 23.14 31.20 3.36 10.78 89
CZ2025 42.84 22.16 29.61 3.30 11.14
BLUP 41.94 25.33 30.41 2.20 7.22
第二节间长度
IL2 (cm)
WJ2025 25.34 12.02 18.89 2.11 11.19 85
CZ2025 20.28 10.32 14.52 1.85 12.77
BLUP 20.89 13.34 16.71 1.19 7.14
第三节间长度
IL3 (cm)
WJ2025 22.34 8.53 13.35 1.89 14.18 75
CZ2025 15.14 5.88 10.03 1.60 15.97
BLUP 15.22 9.79 11.69 0.78 6.67

Fig. 1

Correlation coefficients of first internode length (A), second internode length (B), and third internode length (C) in two different environments and BLUP *, ** and ***, significance at the 0.05, 0.01, and 0.001 probability levels, respectively. The same as below"

Table 2

Coefficients of pairwise Pearson correlations between internode lengths, and agronomic traits"

性状
Trait
第一节间长度
IL1
第二节间长度
IL2
第三节间长度
IL3
株高
PH
有效分蘖数
ETN
穗长
SL
每穗小穗数
SNS
第二节间长度IL2 0.416**
第三节间长度IL3 0.311** 0.748**
株高PH 0.615** 0.556** 0.613**
有效分蘖数ETN 0.033 -0.009 -0.026 -0.098
穗长SL 0.095 -0.04 -0.117 0.283** -0.021
每穗小穗数SNS -0.024 -0.122 -0.042 0.109 -0.188** 0.380**
千粒重TKW 0.04 0.016 -0.031 0.11 0.105 0.198** -0.245**

Fig. 2

Manhattan and quantile-quantile (Q-Q) plots showing the results of genome wide association studies A: IL1; B: IL2; C: IL3. The red dotted line refers to the threshold of -log10(P)=4.00"

Table 3

GWAS-based identification of stably expressed loci for stem internode length"

性状
Trait
位点
QTL
单核苷酸多态性
SNP
等位位点
Allele
染色体
Chromosome
物理位置
Position (bp)
显著性阈值
-log10(P)
贡献率
R2(%)
环境
Environments
第一节间长度
IL1
QIL1.sau.5A 5A_501635166 T/C 5A 501635166 4.21-5.42 1.62-2.19 CZ2025, WJ2025, BLUP
第二节间长度
IL2
QIL2.sau.5A.1 5A_510913304 G/A 5A 510913304 4.23-5.58 5.90-8.18 CZ2025, WJ2025, BLUP
5A_510913376 C/T 5A 510913376 4.21-5.68 5.82-8.30 CZ2025, WJ2025, BLUP
5A_510913381 T/C 5A 510913381 4.21-5.68 5.82-8.30 CZ2025, WJ2025, BLUP
5A_511296202 T/C 5A 511296202 4.02-5.54 5.88-8.59 CZ2025, WJ2025, BLUP
5A_511296204 C/G 5A 511296204 4.02-5.54 5.88-8.59 CZ2025, WJ2025, BLUP
5A_511991250 G/T 5A 511991250 4.01-5.76 5.98-9.16 CZ2025, WJ2025, BLUP
5A_513013678 A/G 5A 513013678 4.02-5.69 5.88-8.88 CZ2025, WJ2025, BLUP
5A_513255742 T/C 5A 513255742 4.04-6.08 5.95-9.65 CZ2025, WJ2025, BLUP
5A_514724687 C/T 5A 514724687 4.14-5.41 8.06-11.04 CZ2025, WJ2025, BLUP
5A_514884600 A/G 5A 514884600 4.04-6.08 5.95-9.65 CZ2025, WJ2025, BLUP
QIL2.sau.5A.2 5A_517883089 G/A 5A 517883089 4.10-6.28 6.10-10.07 CZ2025, WJ2025, BLUP
5A_518500826 T/C 5A 518500826 4.09-6.17 6.08-9.87 CZ2025, WJ2025, BLUP
5A_518579568 A/G 5A 518579568 4.04-6.08 5.95-9.65 CZ2025, WJ2025, BLUP
5A_519456501 C/T 5A 519456501 4.04-6.08 5.95-9.65 CZ2025, WJ2025, BLUP
5A_519549809 G/A 5A 519549809 4.08-5.57 5.69-8.23 CZ2025, WJ2025, BLUP
5A_519551494 C/T 5A 519551494 4.20-5.63 5.90-8.32 CZ2025, WJ2025, BLUP
5A_519604654 T/C 5A 519604654 4.32-6.08 6.09-9.10 CZ2025, WJ2025, BLUP
5A_521267698 G/A 5A 521267698 4.02-5.50 4.55-6.58 CZ2025, WJ2025, BLUP
第三节间长度
IL3
QIL3.sau.4D 4D_20705614 G/A 4D 20705614 4.04-5.03 7.76-10.07 CZ2025, WJ2025, BLUP
4D_21001512 G/T 4D 21001512 4.22-5.28 8.15-10.61 CZ2025, WJ2025, BLUP

Fig. 3

Effects of QIL2.sau.5A.1 (5A_514724687) on agronomic traits"

Fig 4

Effects of QIL2.sau.5A.2 (5A_517883089) on agronomic traits"

Fig. 5

Effects of QIL3.sau.4D (4D_21001512) on agronomic traits"

Table 4

Prediction and annotation of candidate genes for internode lengths in wheat"

位点
QTL
候选基因
Candidate genes
起始位置
Start position (Mb)
基因注释或编码蛋白
Gene annotation or coding proteins
同源基因
Homologs gene
QIL3.sau.4D TraesCS4D02G041400 19.813161 ATP依赖性RNA解旋酶
ATP-dependent RNA helicase
OsRH21, OsRH52B, OsRH52C
TraesCS4D02G042100 20.090627 碱性蓝蛋白
Basic blue protein
OsUCL8, OsUCL9, OsUCL12, ARPN, ENODL22
QIL1.sau.5A TraesCS5A02G286200 494.413105 pfkB样碳水化合物激酶家族蛋白
pfkB-like carbohydrate kinase family protein
OsFKI, OsFKII
QIL2.sau.5A.2 TraesCS5A02G299500 509.602215 组蛋白H4 Histone H4 无Null
TraesCS5A02G299800 509.669584 RNA聚合酶Ⅱ转录辅激活因子
RNA polymerase Ⅱ transcriptional coactivator
SI1, KIWI, KELP
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