# GWAS-Based Mining of Candidate Genes for Low-Nitrogen Tolerance in Maize

**Authors:** Baobao Wang, Luo Xu, Ying Huang, Shaoxin Wang, Zhongjian Li, Rui Guo, Liang Ma, Liping Xu, Zhaohan Yue, Jianying Feng, Dengfeng Zhang

PMC · DOI: 10.3390/ijms27042060 · International Journal of Molecular Sciences · 2026-02-23

## TL;DR

This study identifies genes in maize linked to low-nitrogen tolerance, aiming to improve nitrogen use efficiency for sustainable agriculture.

## Contribution

The study introduces a GWAS-based approach to discover candidate genes and inbred lines for low-nitrogen tolerance in maize.

## Key findings

- 122 significant SNPs in gene regions were detected for low-nitrogen tolerance in maize.
- Three core genes (Zm00001d027880, Zm00001d034047, Zm00001d010574) were identified through GO and KEGG enrichment analyses.
- Inbred lines H1710, 23N272, and 23N41 showed superior low-nitrogen tolerance.

## Abstract

Nitrogen (N) is an essential yield-limiting factor in maize, and identifying genes that improve nitrogen use efficiency (NUE) is critical for sustainable agriculture and environmental protection. However, the genetic basis of NUE in maize remains poorly understood. In this study, we performed a genome-wide association study (GWAS) using a mixed linear model (MLM) controlling for population structure and kinship, based on an association panel of 282 maize inbred lines genotyped via the Maize 50K GBTS array (53,162 SNPs). Ten NUE-related traits (grain yield, hundred-kernel weight, ear length, ear diameter, kernel row number, kernel number per row, SPAD value, ASI, plant height, ear height) were evaluated under two N levels during the 2024–2025 growing seasons. The GWAS analysis detected 122 significant SNPs in gene regions linked to low N tolerance under the studied conditions. Linkage disequilibrium analysis and functional annotation narrowed down 26 candidate genes, whose GO and KEGG enrichment analyses (Fisher’s exact test) identified three core genes (Zm00001d027880, Zm00001d034047, Zm00001d010574). Furthermore, several inbred lines (H1710, 23N272, and 23N41) demonstrating superior low-nitrogen tolerance were identified. The primary subsequent focus in future research for these genetic materials will be their utilization to breed new cultivars with enhanced nitrogen use efficiency.

## Full-text entities

- **Genes:** Diacylglycerol kinase [NCBI Gene 100281194], MADS24 [NCBI Gene 542038], ZmPK1 [NCBI Gene 542378]
- **Diseases:** water deficit (MESH:D000069578), NUE (MESH:D007222), injury to (MESH:D014947), MLM (MESH:D004195), low (MESH:D009800)
- **Chemicals:** Chlorophyll (MESH:D002734), starch (MESH:D013213), MDA (MESH:D008315), carbon (MESH:D002244), N (MESH:D009584), CTAB (MESH:D000077286), phosphatidylinositol (MESH:D010716), P (MESH:D010758), K (MESH:D011188), glycerophospholipid (MESH:D020404), inositol phosphate (MESH:D007295), nitrate (MESH:D009566), H2O2 (MESH:D006861), LN (-), NO (MESH:D009614), membrane lipid (MESH:D008563), chlorate (MESH:D002704), sucrose (MESH:D013395), lipid (MESH:D008055)
- **Species:** Homo sapiens (human, species) [taxon 9606], Zea mays (maize, species) [taxon 4577], Arabidopsis thaliana (mouse-ear cress, species) [taxon 3702]

## Full text

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## Figures

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## References

44 references — full list in the complete paper: https://tomesphere.com/paper/PMC12940573/full.md

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Source: https://tomesphere.com/paper/PMC12940573