生物技术通报 ›› 2026, Vol. 42 ›› Issue (8): 97-105.doi: 10.13560/j.cnki.biotech.bull.1985.2025-1211
宋子慧1,2,3, 薛丽3,4, 王森3,5, 孙建昌6, 韩冰3, 崔迪3, 韩龙植3, 赵正武1(
), 马小定2,3(
)
收稿日期:2025-11-10
出版日期:2026-08-26
发布日期:2026-08-17
通讯作者:
赵正武zhaozhengwu513@sina.com基金资助:
SONG Zi-hui1,2,3, XUE Li3,4, WANG Sen3,5, SUN Jian-chang6, HAN Bing3, CUI Di3, HAN Long-zhi3, ZHAO Zheng-wu1(
), MA Xiao-ding2,3(
)
Received:2025-11-10
Published:2026-08-26
Online:2026-08-17
摘要:
目的 分析与水稻叶枯萎度相关的数量性状位点(QTLs),为筛选水稻耐旱候选基因提供参考。 方法 以300份水稻种质资源为材料,在干旱胁迫下调查叶卷曲指数(leaf rolling index, LRI)和叶枯萎度(leaf wilt degree, LWD),利用全基因组关联分析(genome-wide association study, GWAS)定位耐旱相关数量性状位点。 结果 叶卷曲指数和叶枯萎度表型呈正态分布,通过全基因组关联分析定位到4个与叶枯萎度相关的数量性状位点(quantitative trait loci, QTLs),其中,qLWD4-1在2个调查时期均能被检测到。根据单倍型分析和基因功能注释信息,在qLWD4-1定位区间内筛选到4个耐旱候选基因(LOC_Os04g54360、LOC_Os04g54474、LOC_Os04g54560和LOC_Os04g54570)。 结论 在2个调查时期内均检测到一个稳定表达、与叶枯萎度相关的QTL(qLWD4-1),并在该区间内鉴定到4个耐旱候选基因。
宋子慧, 薛丽, 王森, 孙建昌, 韩冰, 崔迪, 韩龙植, 赵正武, 马小定. 基于全基因组关联分析挖掘水稻抗旱相关基因[J]. 生物技术通报, 2026, 42(8): 97-105.
SONG Zi-hui, XUE Li, WANG Sen, SUN Jian-chang, HAN Bing, CUI Di, HAN Long-zhi, ZHAO Zheng-wu, MA Xiao-ding. Mining of Drought-resistance Related Genes in Rice Based on Genome-wide Association Study[J]. Biotechnology Bulletin, 2026, 42(8): 97-105.
性状 Trait | 调查日期 Investigation date | 富源4 Fuyuan 4 | 群体 Population | |||
|---|---|---|---|---|---|---|
| 均值 Mean | 变异范围 Variation range | 均值 Mean | 标准差 SD | 变异系数 CV (%) | ||
| 叶卷曲指数LRI | 7/6 | 2.91 | 1-7 | 3.27 | 1.61 | 49.2 |
| 8/4 | 3.38 | 1-9 | 3.74 | 2.30 | 61.4 | |
| 叶枯萎度LWD | 7/6 | 3.08 | 1-7 | 3.16 | 1.31 | 41.4 |
| 7/13 | 3.51 | 1-7 | 3.66 | 1.30 | 35.4 | |
| 7/24 | 3.84 | 1-7 | 3.81 | 1.27 | 33.3 | |
| 8/4 | 4.54 | 1-9 | 4.77 | 1.98 | 41.5 | |
表1 干旱胁迫下不同时期LRI和LWD的表现
Table 1 Performance of LRI and LWD at different stages under drought stress
性状 Trait | 调查日期 Investigation date | 富源4 Fuyuan 4 | 群体 Population | |||
|---|---|---|---|---|---|---|
| 均值 Mean | 变异范围 Variation range | 均值 Mean | 标准差 SD | 变异系数 CV (%) | ||
| 叶卷曲指数LRI | 7/6 | 2.91 | 1-7 | 3.27 | 1.61 | 49.2 |
| 8/4 | 3.38 | 1-9 | 3.74 | 2.30 | 61.4 | |
| 叶枯萎度LWD | 7/6 | 3.08 | 1-7 | 3.16 | 1.31 | 41.4 |
| 7/13 | 3.51 | 1-7 | 3.66 | 1.30 | 35.4 | |
| 7/24 | 3.84 | 1-7 | 3.81 | 1.27 | 33.3 | |
| 8/4 | 4.54 | 1-9 | 4.77 | 1.98 | 41.5 | |
图1 干旱胁迫下不同时期LRI和LWD的频率分布a‒b:7月6日(a)和8月4日(b)叶卷曲指数;c‒f:7月6日(c)、7月13日(d)、7月24日(e)和8月4日(f)叶枯萎度
Fig. 1 Frequency distribution of LRI and LWD at different stages under drought stressa-b: Leaf rolling index (LRI) measured on July 6 (a) and August 4 (b). c-f: Leaf wilt degree (LWD) scored on July 6 (c), July 13 (d), July 24 (e), and August 4 (f)
| 性状Trait | 调查日期Investigating date | 位点Locus | 染色体Chromosomes | 物理区间 Physical interval (Mb) | SNP数量 Number of SNPs | 峰值SNP Top SNP | 变异解释率 PVE (%) |
|---|---|---|---|---|---|---|---|
| LWD | 7/13 | qLWD1 | 1 | 15.03‒15.49 | 2 | 15 289 931 | 1.20 |
| qLWD4-1 | 4 | 32.06‒32.66 | 2 | 32 260 706 | 1.10 | ||
| LWD | 7/24 | qLWD4-2 | 4 | 31.79‒32.43 | 5 | 32 033 623 | 1.48 |
| qLWD4-1 | 4 | 32.06‒32.66 | 3 | 32 457 574 | 1.56 | ||
| qLWD4-3 | 4 | 32.47‒33.04 | 3 | 32 843 337 | 1.29 |
表2 LWD性状的QTL定位结果
Table 2 QTL mapping for leaf wilt degree (LWD)
| 性状Trait | 调查日期Investigating date | 位点Locus | 染色体Chromosomes | 物理区间 Physical interval (Mb) | SNP数量 Number of SNPs | 峰值SNP Top SNP | 变异解释率 PVE (%) |
|---|---|---|---|---|---|---|---|
| LWD | 7/13 | qLWD1 | 1 | 15.03‒15.49 | 2 | 15 289 931 | 1.20 |
| qLWD4-1 | 4 | 32.06‒32.66 | 2 | 32 260 706 | 1.10 | ||
| LWD | 7/24 | qLWD4-2 | 4 | 31.79‒32.43 | 5 | 32 033 623 | 1.48 |
| qLWD4-1 | 4 | 32.06‒32.66 | 3 | 32 457 574 | 1.56 | ||
| qLWD4-3 | 4 | 32.47‒33.04 | 3 | 32 843 337 | 1.29 |
图3 水稻叶卷曲指数和叶枯萎度性状GWAS结果a‒b:7月6日(a)和8月4日(b)调查的叶卷曲指数GWAS结果,左为曼哈顿图,右为Q-Q图;c‒f:7月6日(c)、7月13日(d)、7月24日(e)和8月4日(f)调查的叶枯萎度GWAS结果,左为曼哈顿图,右为Q-Q图
Fig. 3 GWAS results for LRI and LWD in ricea‒b: GWAS for LRI measured on July 6 (a) and August 4 (b). The left panels show the Manhattan plots, and the right panels display the quantile-quantile (Q-Q) plots. c‒f: GWAS for LWD measured on four dates: July 6 (c), July 13 (d), July 24 (e), and August 4 (f). The left panels show the Manhattan plots, and the right panels display the quantile-quantile (Q-Q) plots
候选基因 Candidate gene | 基因注释 Gene annotation |
|---|---|
| LOC_Os04g53990 | 乙烯反应蛋白相关 Ethylene-responsive protein related |
| LOC_Os04g54330 | 乙酰转移酶GNAT家族 Acetyltransferase GNAT family |
| LOC_Os04g54360 | 含糖基转移酶8结构域的蛋白质 Glycosyl transferase 8 domain containing protein |
| LOC_Os04g54474 | 转录因子 Transcription factor |
| LOC_Os04g54560 | 转移酶家族蛋白 Transferase family protein |
| LOC_Os04g54570 | 转移酶家族蛋白 Transferase family protein |
| LOC_Os04g54800 | 莽草酸激酶 Shikimate kinase |
| LOC_Os04g54900 | bHLH家族蛋白 bHLH family protein |
表3 qLWD4-1位点的候选基因
Table 3 Candidate genes for the qLWD4-1 locus
候选基因 Candidate gene | 基因注释 Gene annotation |
|---|---|
| LOC_Os04g53990 | 乙烯反应蛋白相关 Ethylene-responsive protein related |
| LOC_Os04g54330 | 乙酰转移酶GNAT家族 Acetyltransferase GNAT family |
| LOC_Os04g54360 | 含糖基转移酶8结构域的蛋白质 Glycosyl transferase 8 domain containing protein |
| LOC_Os04g54474 | 转录因子 Transcription factor |
| LOC_Os04g54560 | 转移酶家族蛋白 Transferase family protein |
| LOC_Os04g54570 | 转移酶家族蛋白 Transferase family protein |
| LOC_Os04g54800 | 莽草酸激酶 Shikimate kinase |
| LOC_Os04g54900 | bHLH家族蛋白 bHLH family protein |
候选基因 Candidate gene | 单倍型 Haplotype | 叶枯萎度 Leaf wilt degree | 基因注释Gene annotation | |
|---|---|---|---|---|
| 7/13 | 7/24 | |||
| LOC_Os04g54360 | Hap_1:ACCTCCGG | - | 3.79b | 含糖基转移酶8结构域的蛋白质 |
| Hap_2:ACCTCCGA | - | 4.01a | Glycosyl transferase 8 domain containing protein, putative, expressed | |
| Hap_3:ACTTCCAG | - | 3.63b | ||
| Hap_4:ACCTTCAG | - | 3.55c | ||
| LOC_Os04g54474 | Hap_1:GGTAGAG | 3.59b | 3.73b | bZIP转录因子 |
| Hap_2:GGTATAG | 3.80a | 4.02a | The bZIP family of transcription factors | |
| LOC_Os04g54560 | Hap_1:CGCCCGCGGCGGACCCGGA | 3.52b | 3.70b | 转移酶家族蛋白 Transferase family protein |
| Hap_2:CACCCGCGGCGGACCCGGA | 3.78a | 4.00a | ||
| Hap_3:CATCCGCGGCGGACCCGGA | 3.60ab | 3.60b | ||
| LOC_Os04g54570 | Hap_1:GGCG | 3.60b | 3.75b | 转移酶家族蛋白 |
| Hap_2:AGCG | 3.80a | 4.04a | Transferase family protein | |
表4 qLWD4-1候选基因的单倍型分析
Table 4 Haplotype analysis of candidate gene qLWD4-1
候选基因 Candidate gene | 单倍型 Haplotype | 叶枯萎度 Leaf wilt degree | 基因注释Gene annotation | |
|---|---|---|---|---|
| 7/13 | 7/24 | |||
| LOC_Os04g54360 | Hap_1:ACCTCCGG | - | 3.79b | 含糖基转移酶8结构域的蛋白质 |
| Hap_2:ACCTCCGA | - | 4.01a | Glycosyl transferase 8 domain containing protein, putative, expressed | |
| Hap_3:ACTTCCAG | - | 3.63b | ||
| Hap_4:ACCTTCAG | - | 3.55c | ||
| LOC_Os04g54474 | Hap_1:GGTAGAG | 3.59b | 3.73b | bZIP转录因子 |
| Hap_2:GGTATAG | 3.80a | 4.02a | The bZIP family of transcription factors | |
| LOC_Os04g54560 | Hap_1:CGCCCGCGGCGGACCCGGA | 3.52b | 3.70b | 转移酶家族蛋白 Transferase family protein |
| Hap_2:CACCCGCGGCGGACCCGGA | 3.78a | 4.00a | ||
| Hap_3:CATCCGCGGCGGACCCGGA | 3.60ab | 3.60b | ||
| LOC_Os04g54570 | Hap_1:GGCG | 3.60b | 3.75b | 转移酶家族蛋白 |
| Hap_2:AGCG | 3.80a | 4.04a | Transferase family protein | |
图4 20% PEG-6000胁迫处理下4个候选基因的表达水平a, c, e, g: 9311. b, d, f, h: Nip. * P<0.05; ** P<0.01; *** P<0.001; **** P<0.000 1
Fig. 4 Expression levels of four candidate genes under 20% PEG-6000 stress treatment
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