Scientia Agricultura Sinica ›› 2026, Vol. 59 ›› Issue (21): 4746-4764.doi: 10.3864/j.issn.0578-1752.2026.21.004

• TILLAGE & CULTIVATION·PHYSIOLOGY & BIOCHEMISTRY·AGRICULTURE INFORMATION TECHNOLOGY • Previous Articles     Next Articles

Effects of Late Spring Coldness at the Anther Connective Stage on the Morphology and Metabolomics of Superior and Inferior Grains in Wheat

WENG Ying(), WANG Hao, LIU ZeHao, GUO YiTing, TANG YaTing, LIU Liu, ZHENG BaoQiang, LI JinCai(), CHEN Xiang()   

  1. College of Agriculture, Anhui Agricultural University/National Engineering Laboratory of Crop Stress Resistance Breeding/Huaibei Field Comprehensive Scientific Observation and Research Station, Ministry of Agriculture and Rural Affairs, Hefei 230036
  • Received:2026-01-05 Accepted:2026-09-20 Online:2026-11-01 Published:2026-11-06
  • Contact: LI JinCai, CHEN Xiang

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Abstract:

【Objective】Late spring coldness is a major agro-meteorological disaster constraint limiting large-scale yield improvement of wheat in the Huang-Huai Plain of China. It not only affects spikelet and floret development but also influences grain weight. This study combined grain phenotype determination and untargeted metabolomics analysis to systematically investigate the morphology, grain weight, and metabolic responses of superior and inferior grains under varying degrees of late spring coldness at the anther connective stage, aiming to elucidate the intrinsic mechanisms underlying the differences in grain filling between superior and inferior grains of wheat with different resistances to late spring coldness from the perspectives of grain sink capacity formation and metabolic regulation. 【Method】In this study, the cold-tolerant wheat cultivar Yannong 19 (YN19) and the cold-sensitive cultivar Xinmai 26 (XM26) were used as experimental materials. Low-temperature treatments were conducted in an artificial climate chamber at the anther connective stage of young spike differentiation to simulate late spring coldness conditions in the field, with three temperature gradients (CK: 10 ℃; T1: 2 ℃; T2: −2 ℃). At maturity, the main agronomic traits of wheat were investigated. Grain morphology and grain weight were dynamically measured at 7, 14, 21, 28, and 35 days after anthesis to analyze the changes in grain sink capacity and grain weight under different temperature treatments. Superior and inferior grains were collected at 14 and 21 days after anthesis, respectively, and untargeted metabolomics analysis was performed using liquid chromatography-mass spectrometry (LC-MS). Multivariate statistical analysis was combined to screen differential metabolites, and the shared and specific responses of metabolites among different cultivars, stress intensities, grain-filling stages, and grain positions were compared. KEGG was used for metabolic pathway enrichment analysis. 【Result】Late spring coldness significantly inhibited the increase in grain sink capacity and reduced grain volume at maturity. In both wheat cultivars, the thousand-grain weight of superior and inferior grains decreased significantly with increasing stress intensity, with a more pronounced reduction observed in inferior grains. Metabolomic analysis identified a total of 2 474 metabolites, including 1 118 annotated metabolites. The number of differentially accumulated metabolites increased markedly with increasing intensity of late spring coldness, and XM26 exhibited more differentially accumulated metabolites than YN19. Only a small number of common differentially accumulated metabolites were shared among different treatments and cultivars. Notably, L-phenylalanine in superior grains at 21 d was identified as a common significantly different metabolite in multiple comparison groups. KEGG pathway enrichment analysis revealed that, under late spring coldness, differentially accumulated metabolites in superior grains were mainly enriched in energy metabolism-related pathways, such as the tricarboxylic acid (TCA) cycle and glycolysis/gluconeogenesis. In contrast, differentially accumulated metabolites in inferior grains were predominantly enriched in pathways associated with energy compensation and antioxidative responses, including starch and sucrose metabolism, oxidative phosphorylation, and glutathione metabolism. 【Conclusion】Late spring coldness exacerbated the differences in grain filling between superior and inferior grains by disrupting energy metabolism and assimilate accumulation during the grain-filling period. These findings provided the metabolomic evidence for elucidating the damage mechanisms of late spring coldness in wheat and for the breeding of cold-tolerant wheat cultivars.

Key words: wheat, late spring coldness, superior and inferior grains, metabolome, differentially accumulated metabolites

Fig. 1

Temperature change in the artificial climate chamber during the experiment CK: Control temperature at 10 ℃; T1: Low-temperature treatment at 2 ℃; T2: Low-temperature treatment at -2 ℃. The same as below"

Table 1

Effect of late spring coldness on the agronomic traits of two wheat varieties"

品种
Variety		 处理
Treatment		 株高
PH (cm)		 重心高度
CGH (cm)		 穗下节间长
SIL (cm)		 穗长
SL (cm)		 穗宽
SW (mm)		 总小穗数
TS		 结实小穗数
FS		 小穗结实率
SSR (%)		 产量
YP (g/pot)
YN19 CK 74.36±0.43a 49.70±0.70a 21.92±0.17a 9.98±0.08b 11.76±0.40b 21.80±0.37ab 19.80±0.37a 90.82±0.15a 32.86±0.39b
T1 71.44±1.45b 46.70±0.75b 20.22±0.66b 9.72±0.09c 11.47±0.26b 21.60±0.51ab 19.40±0.40ab 89.85±0.74a 30.72±0.59cd
T2 68.50±1.27c 44.40±1.03c 18.94±0.71c 9.48±0.08d 10.26±0.16c 21.00±0.45ab 18.60±0.40bc 88.60±1.06ab 29.29±0.42d
XM26 CK 66.22±0.88d 42.54±0.76d 15.90±0.38d 10.46±0.17a 13.43±0.33a 22.00±0.32a 20.00±0.32a 90.90±0.13a 39.71±1.66a
T1 62.44±0.85e 38.06±1.35e 14.30±0.39e 10.00±0.04b 11.40±0.41b 21.20±0.20ab 18.40±0.24c 86.79±0.93bc 32.35±0.94bc
T2 60.80±0.54f 37.22±0.29e 12.56±0.55f 9.72±0.20c 10.44±0.12c 20.60±0.40b 17.40±0.24d 84.55±1.59c 30.26±1.32d
F值
F-value		 F-Variety 412.36** 432.41** 894.33** 32.34** 6.01* 0.38 7.09* 12.27** 40.15**
F-Treatment 64.53** 73.12** 79.88** 37.62** 28.93** 3.82* 19.22** 13.89** 61.06**
F-V×T 0.88 1.77 0.47 1.6 5.05* 0.38 3.05 3.48 13.99**

Fig. 2

Grain size changes of superior and inferior wheat grains at different days after anthesis G1, G2 and G3 denote the first, second and third grain positions within a spikelet, respectively"

Fig. 3

Effects of late spring coldness on the 1000-grain weight of superior and inferior grains at different post-flowering periods in two wheat varieties SG: Superior grain; IG: Inferior grain. Different lowercase letters indicate significant differences among different treatments at the 0.05 probability level"

Fig. 4

Principal component analysis and correlation analysis of metabolites A: PC1 and PC2 indicate the first and second principal components, respectively, and the percentage represents the explanatory power of the principal component on the dataset; B: The vertical and diagonal axes represent the sample names of different samples. For Pearson correlation coefficients greater than 0, the color depth represents the strength of the positive correlation: the deeper the color, the stronger the positive correlation"

Fig. 5

Classification and statistical analysis of metabolites"

Fig. 6

Analysis of the number of differentially accumulated metabolites"

Fig. 7

Venn diagram of differentially accumulated metabolites at 14 and 21 days after anthesis"

Fig. 8

Fold change plot of differentially accumulated metabolites of superior and inferior grains at 14 days after anthesis between two wheat varieties The horizontal coordinate represents the fold change value of differential metabolite expression, and the vertical coordinate represents the names of the metabolites. The same as below"

Fig. 9

Fold change plot of differentially accumulated metabolites of superior and inferior grains at 21 days after anthesis between two wheat varieties"

Fig. 10

Enrichment map of differentially accumulated metabolites of superior and inferior grains at 14 days after anthesis between two wheat varieties Each bubble in the figure represents a pathway, the horizontal coordinate represents Rich factor and the vertical coordinate represents the metabolic pathway name. The colour of the bubble represents the enrichment significance (P value), the redder the colour, the more significantly the pathway is enriched, and the size of the bubble represents the number of differentially accumulated metabolites enriched to the pathway. The first 20 pathways with the smallest P value are selected for display. The same as below"

Fig. 11

Enrichment map of differentially accumulated metabolites of superior and inferior grains at 21 days after anthesis between two wheat varieties"

Fig. 12

Changes in the metabolic network of superior and inferior grains under late spring coldness stress A: Superior grain; B: Inferior grain"

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