核桃无融合生殖核仁内源激素含量变化与基因表达分析

doi:10.3969/j.issn.1004-1524.20231390

浙江农业学报 ›› 2025, Vol. 37 ›› Issue (2): 381-393.DOI: 10.3969/j.issn.1004-1524.20231390

核桃无融合生殖核仁内源激素含量变化与基因表达分析

陈凤(), 陈虹(), 陈兵权, 宝春杰, 周昊亮, 赵鑫, 郭来珍

新疆农业大学林学与风景园林学院,新疆乌鲁木齐 830052

收稿日期:2023-12-12 出版日期:2025-02-25 发布日期:2025-03-20
作者简介:*陈虹,E-mail:ch333999@126.com
陈凤(1999—),女,重庆人,硕士研究生,研究方向为森林培育。E-mail:chen98036@163.com
通讯作者: 陈虹
基金资助:
新疆维吾尔自治区自然科学基金(2022D01A180);国家自然科学基金(31960325);国家自然科学基金(31760219)

Analysis of endogenous hormone changes and gene expression related to walnut apomixis kernels formation

CHEN Feng(), CHEN Hong(), CHEN Bingquan, BAO Chunjie, ZHOU Haoliang, ZHAO Xin, GUO Laizhen

College of Forestry and Landscape Architecture, Xinjiang Agricultural University, Urumqi 830052, China

Received:2023-12-12 Online:2025-02-25 Published:2025-03-20
Contact: CHEN Hong

摘要/Abstract

摘要：

为明确核桃无融合生殖核仁的调控机制,以新新2号核桃(花后60 d、花后80 d、花后100 d)的核仁为试验材料,运用转录组分析无融合生殖和正常授粉受精核仁的差异表达基因(DEG),并测定核仁中的生长素(3-indoleacetic acid, IAA)、细胞分裂素(cytokinin, ZT)、赤霉素(gibberellin, GA)、油菜素甾醇(brassinosteroid, BR)、脱落酸(abscisic acid, ABA)和乙烯(ethylene, ETH)等激素含量变化。研究结果表明:花后60 d,无融合生殖核仁的IAA、ZT、BR含量显著(P<0.05)低于正常授粉受精核仁,分别降低28.57%、19.59%和12.07%;GA、ABA、ETH含量显著(P<0.05)高于正常授粉受精核仁,分别提高34.91%、31.50%和19.54%。花后80 d,无融合生殖核仁的IAA、ZT、ETH含量显著(P<0.05)高于正常授粉受精核仁,分别提高6.58%、17.80%、7.44%;GA、BR、ABA含量显著(P<0.05)低于正常授粉受精核仁,分别降低29.37%、14.99%、10.65%。花后100 d,无融合生殖核仁的IAA、ZT、GA、BR和ETH含量显著(P<0.05)高于正常授粉受精核仁,分别提高18.87%、30.12%、16.96%、3.70%和20.57%;ABA含量显著(P<0.05)低于正常授粉受精核仁,降低44.85%。转录组分析结果显示,植物激素信号转导路径中共筛选出62个调控植物激素的差异表达基因,其中,生长素流入载体基因(AUX1 LAX_2)、B类细胞分裂素响应因子基因(ARR-B1)、脱落酸受体基因(PYL2)、BAK家族基因(BAK1)、磷酸基团转移酶基因(AHP4)在无融合生殖核仁中上调表达,F-box蛋白基因(GID2_SLY1_2)、E3泛素连接酶基因(EBF1/2_1、EBF1/2_2)在无融合生殖核仁中下调表达。实时荧光定量PCR(qRT-PCR)验证结果显示,挑选出的8个基因的相对表达量差异与转录组分析结果一致。核桃油脂转化初期高含量的IAA、ZT、BR可促进无融合生殖核仁发育,ETH在无融合生殖核仁油脂转化中后期起调控作用。植物激素信号转导途径相关基因可调节激素水平变化,AUX1 LAX_2、ARR-B1、PYL2、BAK1、AHP4、GID2_SLY1_2、EBF1/2基因均参与调控核桃无融合生殖核仁的发育。

关键词: 核桃, 无融合生殖, 核仁, 内源激素, 转录组

Abstract:

In order to understand the molecular regulation of apomictic nucleolus development in walnuts, the apomictic kernels and fertilized kernels of ‘Xinxin 2’ walnut were used as the experimental materials in this study. The transcriptome and hormone contents such as indole-3-acetic acid (IAA), cytokinin (ZT), gibberellin (GA), brassinosteroid (BR), abscisic acid (ABA), and ethylene (ETH) were analyzed. The results showed that at 60 days after flowering, the contents of IAA, ZT, and BR in apomictic kernels were significantly (P<0.05) lower than those in normally fertilized kernels, decreasing by 28.57%, 19.59%, and 12.07%, respectively; the contents of GA, ABA, and ETH in apomictic kernels were significantly (P<0.05) higher than those in normally fertilized kernels, increasing by 34.91%, 31.50%, and 19.54%, respectively. At 80 days after flowering, the contents of IAA, ZT, and ETH in apomictic kernels were significantly (P<0.05) higher than those in normally fertilized kernels, increasing by 6.58%, 17.80%, and 7.44%, respectively; meanwhile, the contents of GA, BR, and ABA in apomictic kernels were significantly (P<0.05) lower than those in normally fertilized kernels, decreasing by 29.37%, 14.99%, and 10.65%, respectively. At 100 days after flowering, the contents of IAA, ZT, GA, BR, and ETH in apomictic kernels were significantly (P<0.05) higher than those in normally fertilized kernels, increasing by 18.87%, 30.12%, 16.96%, 3.70%, and 20.57%, respectively; the content of ABA in apomictic kernels was significantly (P<0.05) lower than those in normally fertilized kernels, decreasing by 44.85%. Transcriptome sequencing result showed that 62 differentially expressed genes (DEGs) involved in plant hormone signal transduction pathways were identified. Among them, auxin influx carrier gene (AUX1 LAX_2), B-class cytokinin response factor gene (ARR-B1), abscisic acid receptor gene (PYL2), BAK family gene (BAK1), and phosphotransferase gene (AHP4) were upregulated in apomictic kernels, whereas F-box protein gene (GID2_SLY1_2) and E3 ubiquitin ligase genes (EBF1/2_1, EBF1/2_2) were downregulated. Eight genes that related to differentially metabolites were verified by qRT-PCR and the results were consistent with the finding in transcriptome analysis. It was concluded that during the oil conversion period of walnuts, high contents of IAA, ZT, BR, ETH could promote the development of apomictic kernels. Genes such as AUX1 LAX_2, ARR-B1, PYL2, BAK1, AHP4, GID2_SLY1_2, and EBF1/2 which connect with plant hormone signal transduction pathways were participating in regulating the development of apomictic kernel development.

Key words: walnut, apomixis, kernel, endogenous hormone, transcriptome

中图分类号:

S664.1

陈凤, 陈虹, 陈兵权, 宝春杰, 周昊亮, 赵鑫, 郭来珍. 核桃无融合生殖核仁内源激素含量变化与基因表达分析[J]. 浙江农业学报, 2025, 37(2): 381-393.

CHEN Feng, CHEN Hong, CHEN Bingquan, BAO Chunjie, ZHOU Haoliang, ZHAO Xin, GUO Laizhen. Analysis of endogenous hormone changes and gene expression related to walnut apomixis kernels formation[J]. Acta Agriculturae Zhejiangensis, 2025, 37(2): 381-393.

图/表 10

表1 候选基因的实时荧光定量PCR引物序列

Table 1 Primer sequences of candidate genes used for real-time fluorescence quantitative PCR

基因ID Gene ID	基因名称 Gene name	正向引物序列 Forward primer sequence (5'→3')	反向引物序列 Reverse primer sequence (5'→3')
c61859_g2	LUX1 LAX_2	CGTTACGTCGGGAGGATGAG	ATTGAGCGACGAAAAGGGGT
c60146_g1	GID2_SLY1_2	TGGCTATTGCAAAATCAGGTC	ACGCTCGAGTGTAGTAGCAAG
c66727_g2	ARR-B2	CTCACCCTTGTAGAGCGAGG	TGCAAACTCAGTGCTAAAATCAA
c65820_g2	AHP4	ACCCATCTTTCACGTGTGTG	TGCCTAATTAGCAATTTCCA
c63550_g3	EBF1/2_1	CGTTACGTCGGGAGGATGAG	CGTTACGTCGGGAGGATGAG
c64965_g1	EBF1/2_2	ACCTTCCCTGCTCCTCTCTT	GAGCCTTGTGGAAGCAAACG
c56774_g2	PYL2	TGTAATTGGGGAATGTTGCA	TGGGAGACACAATGATCGAA
c66515_g2	BAK1_2	AACTGTTGCCGGAGCTTTCT	TGGGATAACACCACATGCAGT

图1 新新2号无融合生殖核仁与正常授粉受精核仁在不同发育时期的内源激素含量无相同字母的表示差异显著(P<0.05)。

Fig.1 The endogenous hormone content of the apomictic kernels and fertilized kernels of ‘Xinxin 2’ walnut at different developmental stages Different letters indicate significant difference at P<0.05.

表2 测序数据质量评估统计表

Table 2 Statistical table for quality assessment of sequencing data

样品编号 Sample ID	原始数据 Raw read	过滤后数据 Clean read	高质量碱基 Clean base/Gb	碱基质量值Quality score		整体测序错误率 Error rate/%	GC含量 GC content/%
样品编号 Sample ID	原始数据 Raw read	过滤后数据 Clean read	高质量碱基 Clean base/Gb	Q20/%	Q30/%	整体测序错误率 Error rate/%	GC含量 GC content/%
G1	52981375	50634989	7.60	95.63	90.14	0.02	45.53
G2	51860163	49469256	7.42	95.38	89.66	0.02	48.78
G3	53160610	50416068	7.56	95.85	90.36	0.02	48.96
W1	45992618	43736529	6.56	94.85	88.63	0.02	44.79
W2	48272843	44792770	6.72	94.87	88.63	0.02	46.45
W3	47683476	44944840	6.74	95.23	89.46	0.02	45.16

图2 新新2号无融合生殖核仁与正常授粉受精核仁中IAA代谢和信号转导相关基因分析 G1~G3和W1~W3分别表示新新2号正常授粉受精核仁与无融合生殖核仁的3个时期。下同。

Fig.2 Analysis of genes related to IAA metabolism and signal transduction in the apomictic kernels and fertilized kernels of ‘Xinxin 2’ walnut G1-G3 and W1-W3 represent the three periods of fertilized kemels and apomictic kemels of ‘Xinxia 2’ walnut.The same as below.

图3 新新2号无融合生殖核仁与正常授粉受精核仁中细胞分裂素代谢和信号转导相关基因分析

Fig.3 Analysis of genes related to cytokinin metabolism and signal transduction in apomictic kernels and fertilized kernels of ‘Xinxin 2’ walnut

图4 新新2号无融合生殖核仁与正常授粉受精核仁中赤霉素代谢和信号转导相关基因分析

Fig.4 Analysis of genes related to gibberellin metabolism and signal transduction in apomictic kernels and fertilized kernels of ‘Xinxin 2’ walnut

图5 新新2号无融合生殖核仁与正常授粉受精核仁中乙烯代谢和信号转导相关基因分析

Fig.5 Analysis of genes related to ethylene metabolism and signal transduction in apomictic kernels and fertilized kernels of ‘Xinxin 2’ walnut

图6 新新2号无融合生殖核仁与正常授粉受精核仁中脱落酸代谢和信号转导相关基因分析

Fig.6 Analysis of genes related to abscisic acid metabolism and signal transduction in apomictic kernels and fertilized kernels of ‘Xinxin 2’ walnut

图7 新新2号无融合生殖核仁与正常授粉受精核仁中油菜素甾醇代谢和信号转导相关基因分析

Fig.7 Analysis of genes related to brassinosteroid metabolism and signal transduction in apomictic kernels and fertilized kernels of ‘Xinxin 2’ walnut

图8 差异表达基因的相对表达量

Fig.8 Relative expression of differentially expressed genes

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核桃无融合生殖核仁内源激素含量变化与基因表达分析

Analysis of endogenous hormone changes and gene expression related to walnut apomixis kernels formation

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