Genome-Wide Association Study of Okra Seed Germination Rate under Seawater
Stress Conditions

doi:10.3969/j.issn.1009-7791.2026.01.002

Subtropical Plant Science ›› 2026, Vol. 55 ›› Issue (1): 12-26.DOI: 10.3969/j.issn.1009-7791.2026.01.002

• Research articles • Previous Articles Next Articles

Genome-Wide Association Study of Okra Seed Germination Rate under Seawater Stress Conditions

HUA Xin-tao^#, TANG Zi-qi^#, YANG Yun, ZHANG Yan, LI Hui-wen, LI Yi-ying, SHU Yang-pei, SUN Jian^*, XIE Dong-wei^*

(School of Life Sciences, Nantong University, Nantong 226000, Jiangsu China)

Received:2025-09-04 Accepted:2025-09-29 Online:2026-02-28 Published:2026-02-28
Contact: SUN Jian; XIE Dong-wei

海水胁迫下黄秋葵种子发芽率的全基因组关联分析

华鑫涛^#，汤子祺^#，杨芸，张妍，李慧雯，李奕莹，束杨培，孙健^*，谢冬微^*

(南通大学生命科学学院，江苏南通 226000)

通讯作者: 孙健；谢冬微
基金资助:
南通市社会民生科技计划项目(MS22021029)；大学生创新创业训练计划项目(202410304109Y)；南通大学大型仪器开放基金项目(KFJN2554)

Abstract

Abstract: To explore the candidate genes related to salt tolerance in okra under seawater stress, 180 okra (Abelmoschus esculentus) germplasm resources from various sources were used as materials, and the seeds were treated with seawater at 3‰ and 6‰ salinity. The germination percentage and relative germination percentage were measured, genome-wide association study (GWAS) was performed with SNP markers, and candidate gene screening was performed by transcriptome sequencing (RNA-seq). The results showed that 3‰ and 6‰ salinity of seawater had different degrees of inhibition on the germination rate of okra seeds. A total of 75 SNP loci were screened out by GWAS. An association interval was detected on chromosome 54, and four SNP loci were detected simultaneously under seawater stress of 3‰ and 6‰ salinity, which were important loci for controlling the relative germination rate and salt tolerance, and a total of 54 annotated genes were detected within 100 kb of the upstream and downstream of the four SNPs. Based on the results of transcriptome sequencing under seawater stress at the seedling stage of okra, six differentially expressed genes were screened out as candidate genes for salt tolerance of okra under seawater stress, namely Ae54G020550, Ae54G020570, Ae54G020630, Ae54G020730, Ae54G020810 and Ae54G020890, and the annotation functions were cytoskeletal composition, encoding CYSTM protein, NPH3 or BTB/POZ protein, alcohol dehydrogenase, leucine-rich repeats, and PPR family proteins. These results provide a reference for analyzing the function of salt tolerance genes in okra and the molecular mechanism of salt tolerance.

Key words: okra, seawater stress, genome-wide association study, transcriptome, candidate gene

摘要： 以不同来源的180份黄秋葵Abelmoschus esculentus种质资源为材料，对其种子进行3‰和6‰盐度海水处理，测定发芽率及相对发芽率，并与SNP标记进行全基因组关联分析(GWAS)，结合转录组测序(RNA-seq)筛选耐盐候选基因。结果表明，3‰和6‰盐度海水对黄秋葵种子发芽率均产生不同程度的抑制作用。GWAS共筛选出75个与耐盐性状显著相关的SNP位点。在54号染色体上检测到一个关联区间，该区间有4个SNP位点在3‰和6‰盐度海水胁迫下均被同时检测到，是控制相对发芽率耐盐性的重要位点，4个SNP上下游100 kb范围内共有54个注释基因。利用黄秋葵苗期海水胁迫转录组测序数据，筛选出6个差异表达基因作为海水胁迫下黄秋葵耐盐性的候选基因，分别为Ae54G020550、Ae54G020570、Ae54G020630、Ae54G020730、Ae54G020810和Ae54G020890，注释功能分别为细胞骨架组成结构、编码CYSTM蛋白、NPH3或BTB/POZ蛋白、醇脱氢酶、富含亮氨酸的重复序列及PPR家族蛋白。研究结果为分析黄秋葵耐盐基因的功能及其耐盐性分子机制提供参考。

关键词: 黄秋葵, 海水胁迫, 全基因组关联分析, 转录组, 候选基因

CLC Number:

Q943

HUA Xin-tao, TANG Zi-qi, YANG Yun, ZHANG Yan, LI Hui-wen, LI Yi-ying, SHU Yang-pei, SUN Jian, XIE Dong-wei. Genome-Wide Association Study of Okra Seed Germination Rate under Seawater Stress Conditions[J]. Subtropical Plant Science, 2026, 55(1): 12-26.

华鑫涛, 汤子祺, 杨芸, 张妍, 李慧雯, 李奕莹, 束杨培, 孙健, 谢冬微. 海水胁迫下黄秋葵种子发芽率的全基因组关联分析[J]. 亚热带植物科学, 2026, 55(1): 12-26.

References

[1] Food and Agriculture Organization of the United Nations. Global status of salt-affected soils – Main report [R]. Rome: FAO, 2024.

[2] 曹帮华, 曹曦东, 刘业仕, 周亮. 我国滨海盐碱地改良及其应用潜力[J]. 中国农民合作社, 2025(6): 21–24.

[3] 杨文涛, 王琦, 郭二丹, 徐皓. 土壤盐渍化对植物影响的研究进展[J]. 农业与技术, 2024, 44(18): 95–99.

[4] 李飞飞, 刘杨, 徐飞. 种子萌发的奥秘: 生理、生态与分子的综合解析[J]. 生命科学, 2024, 36(12): 1470–1477.

[5] 裴云霞, 倪天虹, 乔子瑜, 许辉, 张强. 盐胁迫对结球甘蓝种子萌发、幼苗生长及其生理特性的影响[J/OL]. 云南农业大学学报(自然科学), 2025: 1–8. https://link.cnki.net/urlid/53.1044.S. 20250812.0957.001.

[6] 汪艳, 师尚礼, 于海涛, 韩宜霖. 18个紫花苜蓿品种萌发期耐盐性评价[J/OL]. 草地学报, 2025: 1–17. https://link.cnki.net/ urlid/11.3362.S.20250821.1753.006.

[7] 于婵, 张依琳, 李秋莹, 周艳, 李贺敏, 夏至, 黄勇. 盐碱胁迫对牛至种子萌发和幼苗生理生化特性的影响[J]. 草地学报, 2024, 32(6): 1882–1892.

[8] Naveen K A, Tahmish F, Jitendra M, Isha M, Sushma V, Renu V, Maya V, Ankita B, Priyanka V, Priya M, Chanda B. Halo-tolerant plant growth promoting rhizobacteria for improving productivity and remediation of saline soils [J]. Journal of Advanced Research, 2020(26), 69–82.

[9] 林加发. 海水胁迫对5种豆科植物种子萌发的影响[J]. 浙江农业科学, 2021, 62(10): 2005–2007.

[10] 苏彦宾, 刘鲁江, 亓德明, 郭玲娟, 曾祥伟, 王富强, 张天柱. 海水胁迫对2种景天种子萌发及幼苗生长的影响[J]. 中国农学通报, 2017, 33(3): 88–93.

[11] 赵作章, 陈劲松, 彭尔瑞, 李荣亮, 吴大真. 土壤盐渍化及治理研究进展[J]. 中国农村水利水电, 2023(6): 202–208.

[12] 陈思蓉, 李晨, 孙炳蕊. 水稻耐盐分子机制研究进展[J]. 广东农业科学, 2023, 50(12): 29–42.

[13] 曹英杰, 杨剑飞, 王宇. 全基因组关联分析在作物育种研究中的应用[J]. 核农学报, 2019, 33(8): 1508–1518.

[14] 冯建英, 温阳俊, 张瑾, 章元明. 植物关联分析方法的研究进展[J]. 作物学报, 2016, 42(7): 945–956.

[15] 黄紫红, 范世航, 华玮, 付正伟. 甘蓝型油菜耐盐性全基因组关联分析与种质筛选[J/OL]. 中国油料作物学报, 2025: 1–12. https://doi.org/10.19802/j.issn.1007–9084.2024302.

[16] 秦莉, 马庆, 路小铎. 玉米萌发期耐盐性全基因组关联分析与耐盐候选基因鉴定[J]. 玉米科学, 2025, 33(2): 21–27.

[17] 秦慧彬, 任觐, 娄坤, 商春悦, 阳世杰, 侯森, 李萌, 王海岗, 穆志新. 山西谷子萌发期耐盐性鉴定及相关性状GWAS分析[J]. 植物遗传资源学报, 2025, 26(1): 108–119.

[18] 黄阿根, 陈学好, 高云中, 车婧. 黄秋葵的成分测定与分析[J]. 食品科学, 2007(10): 451–455.

[19] 王君耀, 周峻, 汤谷平. 黄秋葵抗疲劳作用的研究[J]. 中国现代应用药学, 2003(4): 316–317.

[20] 王璐, 李蓓蓓, 王参政, 方柳影, 刘苗苗, 李彦灵. 秋葵果胶对糖尿病小鼠降糖、降脂作用研究[J]. 中国食品添加剂, 2023, 34(9): 236–243.

[21] 谭亮, 杲秀珍, 云忠祥, 俞泽帆, 汪园, 王环. 黄秋葵抗氧化活性研究[J]. 中成药, 2023, 45(3): 829–835.

[22] Md A S, Emon M, Ali M W H, Md Sakib A H, Md Arif H, Imam H R, Moushumi A M, Mst. Sumaia A, Sujoy D, Muhammad T. Anticancer activity of Abelmoschus esculentus (L.) Moench against lung cancer cell: A literature review [J]. Pharmacological Research – Natural Products, 2025, 8: 100317.

[23] 王永慧, 陈建平, 张培通, 高进, 周春霖, 施庆华, 周汝琴. 盐胁迫对不同基因型黄秋葵苗期生长及生理生态特征的影响[J]. 华北农学报, 2016, 31(6): 105–110.

[24] 李啟萌, 宗宪春. 盐胁迫对红黄两种秋葵种子萌发的影响[J]. 中国林副特产, 2021(2): 4–6.

[25] 戴志刚, 康泽培, 谢冬微, 刘岩，孙健. 海水胁迫下黄秋葵种质资源苗期耐盐性鉴定及综合评价[J]. 种子, 2022, 41(8): 42–48.

[26] 刘畅, 程玉静, 王小秋, 仇亮, 刘岩, 戴志刚, 谢冬微, 孙健. 海水胁迫下外源海藻糖对黄秋葵幼苗耐盐性的影响[J]. 中国瓜菜, 2023, 36(8): 84–91.

[27] 唐宁, 张边江, 王立科, 杨平, 陈全战. 海水胁迫对黄秋葵光合及抗氧化生理特性的影响[J]. 分子植物育种, 2019, 17(8): 2639–2645.

[28] 练冬梅, 赖正锋, 姚运法, 洪建基. 黄秋葵盐胁迫下的转录组分析[J]. 中国蔬菜, 2021(6): 42–48.

[29] 曹亚萍, 陈昭昱, 王勇飞, 贾孟君, 贺嘉欣, 张鑫瑞, 李政, 乔永刚, 宋芸. Na₂CO₃对黄秋葵种子萌发及生理特性的影响[J]. 种子, 2020, 39(3): 24–28.

[30] Mckenne A, Hanna M, Banks E, Sivachenko A, Cibulskis K, Kernytsky A, Garimella K, Altshuler D, GABRIEL S, Daly M, Depristo M A. The genome analysis toolkit: a MapReduce framework for analyzing next-generation DNA sequencing data [J]. Genome Research, 2010, 20(9), 1297–1303.

[31] Cingolani P, Platts A, Wang L L, Coon M, Nguyen T, Wang L, Land S J, Lu X Y, Ruden D M. A program for annotating and predicting the effects of single nucleotide polymorphisms, SnpEff: SNPs in the genome of Drosophila melanogaster strain w1118; iso–2; iso–3 [J]. Fly(Austin), 2012, 6(2): 80–92.

[32] Alexander D H, Novembre J, Lange K. Fast model-based estimation of ancestry in unrelated individuals [J]. Genome Research, 2009, 19(9): 1655–1664.

[33] Price A, Patterson N J, Plenge R M, Weinblatt M E, Shadick N A, Reich D. Principal components analysis corrects for stratification in genome-wide association studies [J]. Nature Genetics, 2006, 38(8): 904–909.

[34] Kang H M, Sul J H, Service S K, Zaitlen N A, Kong S Y, Freimer N B, Saba T C, Eekin E. Variance component model to account for sample structure in genome-wide association studies [J]. Nature Genetics, 2010, 42(4): 348–354.

[35] Ginestet C E. ggplot2: Elegant graphics for data analysis [J]. Journal of The Royal Statistical Society Series A–statistics in Society, 2011, 174: 245–246.

[36] Turner S D. qqman: an R package for visualizing GWAS results using Q-Q and manhattan plots [J]. Journal of Open Source Software, 2018, 3(25): 731.

[37] 刘岩，程玉静，翟彩娇，王小秋，刘畅，谢冬微，孙健，戴志刚. 海水胁迫下黄秋葵Na⁺、K⁺含量变化规律研究[J]. 植物学研究, 2021, 10(4): 576–586.

[38] Zhan Y H, Wu T T, Zhao X, Wang Z Q, Chen Y. Comparative physiological and full-length transcriptome analyses reveal the molecular mechanism of melatonin-mediated salt tolerance in okra (Abelmoschus esculentus L.) [J]. BMC Plant Biology, 2021, 21(1): 180.

[39] Livak K J, Schmittgen T D. Analysis of relative gene expression data using real-time quantitative PCR and the 2^–ΔΔCt method [J]. Methods, 2001, 25(4): 402–408.

[40] 汪斌强, 马大卫, 马彦军. 盐胁迫对万寿菊种子萌发和幼苗生长的影响[J]. 甘肃农业大学学报, 2025, 60(3): 189–195.

[41] 李岩岩. 盐胁迫对苦参种子萌发和幼苗生长的影响[J]. 新农民, 2025(3): 63–65.

[42] 丁燕, 呼凤兰, 畅博奇. NaCl胁迫对玉米种子萌发特性及α-淀粉酶活性的影响[J]. 黑龙江农业科学, 2019(4): 11–14.

[43] 曹阳, 张奥, 张传领, 贾鑫, 薛焱, 王晓琴. 蒙古黄芪种子萌发和幼苗生长对盐碱胁迫的响应[J]. 种子, 2023, 42(11): 101–105.

[44] 邓焕莹, 钱峰, 周琬敏, 陈雨寒, 颜统利, 赵燕昊, 蒋玉蓉, 陆国权. 盐胁迫对藜麦种子萌发生长的影响及耐盐机理研究[J]. 福建农业学报, 2023, 38(10): 1139–1145.

[45] 朱蕾, 王向锋, 傅缨, 毛同林. 植物细胞骨架与气孔运动[J]. 科学通报, 2025, 70(25): 4350–4359.

[46] Kumar S, Jeevaraj T, Yunus M H, Chakraborty S, Chakraborty N. The plant cytoskeleton takes center stage in abiotic stress responses and resilience [J]. Plant, Cell and Environment, 2023, 46: 5–22.

[47] Xu Y, Yu Z, Zhang S, Wu C, Yang G, Yan K, Zheng C, Huang J. CYSTM3 negatively regulates salt stress tolerance in Arabidopsis [J]. Plant Molecular Biology, 2019, 99(4–5): 395–406.

[48] Sullivan S, Waksman T, Paliogianni D, Henderson L, Lütkemeyer M, Suetsugu N, Christie J M. Regulation of plant phototropic growth by NPH3/RPT2-like substrate phosphorylation and 14-3-3 binding [J]. Nature Communications, 2021, 12(1): 6129.

[49] 吕彤彤, 颜文慧, 梁艳, 丁寅, 颜庆霞, 李金华. BTB蛋白泛素化介导植物发育和逆境应答的研究进展[J]. 生物工程学报, 2024, 40(1): 63–80.

[50] Ullah U, Mao W, Abbas W, Alharthi B, Bhanbhro N, Xiong M, Gul N, Shalmani A. OsMBTB32, a MATH-BTB domain- containing protein that interacts with OsCUL1s to regulate salt tolerance in rice [J]. Functional & Integrative Genomics, 2023, 23(2): 139.

[51] 沙向红. 盐胁迫对玉米幼苗根系乙醇脱氢酶与乙醛脱氢酶表达的影响[J]. 贵州农业科学, 2015(3): 47–50.

[52] Hao Z Q, Wang T Y, Chen D D, Shen L, Zhang G H, Qian Q, Zhu L. Leucine-rich repeat protein family regulates stress tolerance and development in plants [J]. Rice Science, 2025, 32(1): 32–43.

[53] 李程, 路凯, 王才林, 张亚东. PPR蛋白响应植物非生物胁迫的研究进展[J]. 中国农业科学, 2023, 56(24): 4801–4813.

[54] 吕宝莲, 杨宇昕, 崔立操, 史峰, 马亮, 孔秀英, 张立超, 倪志勇. 小麦bHLH家族转录因子的鉴定及其在盐胁迫条件下的表达分析[J]. 作物杂志, 2024(1): 65–72.

[55] 牛银亭, 鞠乐, 阴志刚, 刘瑞, 曹双, 陈培育, 李君霞. 盐胁迫下谷子叶片转录组de novo测序及分析[J]. 江苏农业科学, 2025, 53(14): 55–62.

[56] 李德珍, 德英, 穆怀彬, 闫伟红, 胡哲华. 盐胁迫下老芒麦叶片转录组SSR位点信息分析[J]. 中国草地学报, 2025, 47(6): 145–150.

Metrics

Copyright © Editorial office of Subtropical Plant Science
Tel: 0592-5654157　E-mail: yrdzwkx@126.com
Supported by Beijing Magtech Co., Ltd.

Genome-Wide Association Study of Okra Seed Germination Rate under Seawater Stress Conditions

海水胁迫下黄秋葵种子发芽率的全基因组关联分析

PDF

Knowledge

Abstract

Cite this article

share this article

References

Related Articles 4

Recommended Articles

Metrics

[1]	ZHONG Feng-di, LING Teng-hong, MOU Gui-ping, ZHU Kong-yu, WANG Peng-long, YAN Xiao-dong. Full-length Transcriptome Sequencing Analysis of Medicinal Plant Pithecellobium clypearia [J]. Subtropical Plant Science, 2024, 53(5): 416-424.
[2]	HONG Yu-yan, YANG Nan, PAN Meng-ting, LIN Mian-fa, XU Wen, XU Hui-long , FAN Shi-ming. Effects of Red and Blue Light Supplementation on Photosynthesis of Tetrastigma hemsleyanum and Its Mechanism [J]. Subtropical Plant Science, 2024, 53(5): 408-415.
[3]	ZHENG Ting-ting,WEI Wei-lin,YANG Xiang-hui,LIN Shun-quan. Development of SSR Molecular Markers Based on Transcriptome Sequencing of Eriobotrya japonica [J]. , 2015, 44(04): 274-278.
[4]	LI Hong. QTL Mapping and Cloning in Plant Genetic Breeding of Disease Resistance [J]. Subtropical Plant Science, 2011, 40(03): 87-92.