甘蓝型冬油菜越冬期矮生长点形态建成及多激素调控机制

doi:10.3864/j.issn.0578-1752.2026.05.003

中国农业科学 ›› 2026, Vol. 59 ›› Issue (5): 951-966.doi: 10.3864/j.issn.0578-1752.2026.05.003

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

甘蓝型冬油菜越冬期矮生长点形态建成及多激素调控机制

刘海卿¹^,²(), 金姣姣¹^,², 孙万仓³, 柴鹏⁴, 祁伟亮¹^,², 杨刚³, 李婵¹, 骆雪梅¹, 苏芸芸¹^,², 秦雪雪¹

¹ 陇东学院农业与生物工程学院，甘肃庆阳 745000
² 甘肃省陇东生物资源保护利用与生态修复重点实验室，甘肃庆阳 745000
³ 甘肃农业大学农学院，兰州 730070
⁴ 甘肃省庆阳市西峰区农业技术推广中心，甘肃庆阳 745000

收稿日期:2025-08-19 接受日期:2025-11-12 出版日期:2026-03-01 发布日期:2026-03-06
联系方式: 刘海卿，E-mail：15117162792@163.com
基金资助:
甘肃省青年科技基金(24JRRM025); 甘肃省农业农村厅种业攻关项目(GYGG-2024-5); 陇东学院青年博士基金(XYBYZK2209); 国家自然科学基金(32360455); 甘肃省高校教师创新基金(甘教技函〔2024〕9号); 甘肃省自然科学基金(22JR5RM205); 庆阳市青年科技人才项目(2025RK1030)

Morphogenesis of the Low-Growth Point and Its Multi-Hormonal Regulatory Mechanism During Overwintering in Winter Rapeseed (Brassica napus L.)

LIU HaiQing¹^,²(), JIN JiaoJiao¹^,², SUN WanCang³, CHAI Peng⁴, QI WeiLiang¹^,², YANG Gang³, LI Chan¹, LUO XueMei¹, SU YunYun¹^,², QIN XueXue¹

¹ School of Agriculture and Bioengineering, Longdong Univerisity, Qingyang 745000, Gansu
² Gansu Key Laboratory of Protection and Utilization for Biological Resources and Ecological Restoration, Qingyang 745000, Gansu
³ College of Agronomy, Gansu Agricultural University, Lanzhou 730070
⁴ Xifeng Agricultural Technology Extension Center, Qingyang 745000, Gansu

Received:2025-08-19 Accepted:2025-11-12 Published:2026-03-01 Online:2026-03-06

摘要/Abstract

摘要：

【目的】 分析植物激素对生长点形态发育及响应抗寒性的作用及机制，以期为甘蓝型冬油菜品种的选育、扩大甘蓝型冬油菜的种植面积及提高单产提供一定的理论依据。【方法】 以9份不同冬性甘蓝型冬油菜品种为材料，在甘肃庆阳连续2年（2023—2025）田间统计越冬率，11月中旬采样进行根/冠、生长点高度等表型鉴定，并结合显微观察（采用石蜡切片和台盼蓝/NBT染色观察生长锥及细胞死亡、超氧阴离子）鉴定不同品种的抗寒性；选取抗寒性由强到弱差异明显的NTS309、天油14号、秦早1号的生长点组织同步进行LC-MS/MS靶向激素组测定和转录组分析，以PCA、O2PLS整合代谢与转录数据，系统解析生长点高度差异的生理与分子基础。【结果】 不同品种越冬率、根颈直径、干重、根/冠、生长点高度差异明显，强抗寒品种低温胁迫后ROS积累少、死亡细胞少，受低温伤害细胞比例小，反之亦然。其中，越冬率与生长点高度呈极显著负相关（r=-0.972，P<0.001）。越冬率>70%的强抗寒品种（NTS309等）生长点高度较低（约3 cm），生长锥高度仅161.5 μm，而弱抗寒品种（秦早1号等）生长点凸出于地表（约5 cm），生长锥高度约252.9 μm。经激素组和转录组联合分析，生长点高度受生长素（Auxin/IAA）-细胞分裂素（CKs）-赤霉素（GA）-脱落酸（ABA）-茉莉酸（JA）等多维网络调控，强抗寒品种游离IAA、结合态IAA、JA-Ile、GA19、ABA-GE显著上调，而iP9G、mT9G、BAP9G下调；强、弱抗寒品种之间的差异表达基因富集于“色氨酸代谢”“玉米素合成”“植物激素信号转导”“α-亚麻酸代谢”通路；O2PLS模型揭示IAA-Glu、tZR、JA-Ile等代谢物与转录模块高度共变，构成激素-转录因子-代谢物反馈环，抑制细胞伸长、促进分生组织休眠，降低生长锥和生长点高度，防御低温伤害。【结论】 明确了甘蓝型冬油菜“矮生长点”是越冬存活的关键形态标记，其形成由多激素网络精细调控。

关键词: 甘蓝型冬油菜, 植物激素, 生长点, 生长锥, 抗寒性

Abstract:

【Objective】 This study aims to clarify the mechanism of plant hormones on the morphology of growth point and their response to cold resistance of Brassica napus, to provide a theoretical basis for breeding winter Brassica napus varieties, expanding its planting area, and increasing yield per unit.【Method】 Using 9 rapeseed varieties with distinct winter hardiness as experimental materials, a two year (2023-2025) consecutive field identification of overwintering rate was conducted in Qingyang, Gansu Province. Mid-November samplings were used to quantify shoot to root ratio, growth point height and additional phenotypic indicators of overwintering potential. Paraffin sectioning and trypan-blue/nitroblue tetrazolium staining were used to visualize growth cone morphology, cell death, and superoxide accumulation to evaluate the cold tolerance of different cultivars. Growth point of three representative genotypes that span the full range of cold hardiness (NTS309>Tianyou 14>Qinzao 1) were simultaneously analysed by targeted LC-MS/MS targeted phytohormone profiling and RNA-seq. PCA and O2PLS integrative analyses of the metabolome and transcriptome, the physiological and molecular mechanisms underlying the differences in growth point height among these varieties were systematically analyzed.【Result】 The results showed the following: Significant differences were observed among cultivars in overwintering rate, root collar diameter, dry weight, root/shoot ratio, and height of the growing point. Strong cold-tolerant varieties, less reactive oxygen species (ROS) and fewer dead cells observed following cold stress, accompanied by a lower proportion of cells damaged by low temperature, the opposite was true for weak cold-tolerant varieties. Statistical analysis revealed that the overwintering rate was extremely significantly and negatively correlated with growth point height (correlation coefficient r=-0.972, P<0.001). Rapeseed varieties with strong cold-tolerance (e.g., NTS309) which exhibited an overwintering rate exceeding 70%, had a relatively lower growth point height (approximately 3 cm), additionally, the height of the growth cone (observed in paraffin sections) was only 161.5 μm. In contrast, varieties with weak cold-tolerance (e.g., Qinzao 1) had growth points that protruded above the soil surface (approximately 5 cm), and their growth cone measured approximately 252.9 μm in height. Integrated analysis of hormone metabolomics and transcriptomics data revealed that the height of rapeseed growth point is regulated by a multi-dimensional regulatory network involving the phytohormones IAA, CKs, GA, JA and ABA. In strong cold-tolerant varieties, the levels of free IAA, conjugated IAA, JA-Ile, GA19, and ABA-GE were significantly up-regulated, whereas those of iP9G, mT9G, and BAP9G were significantly down-regulated. Differentially expressed genes (DEGs) between strong and weak cold-tolerant varieties were significantly enriched in pathways including “tryptophan metabolism”, “zeatin biosynthesis”, “plant hormone signal transduction”, and “α-linolenic acid metabolism”. The orthogonal partial least squares (O2PLS) model indicated that metabolites including IAA-Glu, tZR, and JA-Ile exhibited strong covariance with transcriptional modules, these interactions collectively form a hormone-transcription factor-metabolite feedback loop, which exerts three key regulatory effects: inhibiting cell elongation, promoting meristem dormancy and reducing the height of growth cone and growth points. Ultimately, this regulatory loop enhances the defense capacity of rapeseed against low-temperature freezing damage.【Conclusion】 This study is the first to confirm that the “dwarf growth point” of rapeseed is a key morphological marker for overwintering survival, and its formation is finely regulated by a multi-hormone network.

Key words: winter rapeseed (Brassica napus L.), plant hormone, growth point, growth cone, cold resistance

刘海卿, 金姣姣, 孙万仓, 柴鹏, 祁伟亮, 杨刚, 李婵, 骆雪梅, 苏芸芸, 秦雪雪. 甘蓝型冬油菜越冬期矮生长点形态建成及多激素调控机制[J]. 中国农业科学, 2026, 59(5): 951-966.

LIU HaiQing, JIN JiaoJiao, SUN WanCang, CHAI Peng, QI WeiLiang, YANG Gang, LI Chan, LUO XueMei, SU YunYun, QIN XueXue. Morphogenesis of the Low-Growth Point and Its Multi-Hormonal Regulatory Mechanism During Overwintering in Winter Rapeseed (Brassica napus L.)[J]. Scientia Agricultura Sinica, 2026, 59(5): 951-966.

图/表 8

表1

图1

表2

图2

图3

图4

图5

表3

不同甘蓝型冬油菜品种激素含量分析"

缩略词Index	化合物 Compounds	类别 Class	NTS309	天油14号 Tianyou 14	秦早1号 Qinzao 1	P值 P value
IAA	吲哚-3-乙酸Indole-3-acetic acid	生长素Auxin	24.37±0.56a	17.49±0.19b	28.34±3.34a	0.021
IAA-Glu	吲哚乙酸-谷氨酸Indole-3-acetyl glutamic acid	生长素Auxin	1.79±0.17a	0.38±0.04b	1.58±0.15a	<0.001
MEIAA	吲哚-3-乙酸甲酯Methyl indole-3-acetate	生长Auxin	285.43±30.95a	82.37±14.75b	276.44±18.63a	0.001
IAM	3-吲哚乙酰胺3-Indole acetamide	生长素Auxin	85.23±9.41a	22.91 ±2.23c	55.20±5.32b	0.001
IAN	吲哚-3-乙腈3-Indoleacetonitrile	生长素Auxin	1796.08±69.00a	611.95±67.05b	1597.89±93.47a	<0.001
ICA	吲哚-3-甲酸Indole-3-carboxylic acid	生长素Auxin	45.63±2.15a	22.72±0.56b	22.11±0.48b	<0.001
ICAld	吲哚-3-甲醛Indole-3-carboxaldehyde	生长素Auxin	156.42±24.31a	59.26 ±4.07b	108.93±8.58ab	0.011
IPA	3-吲哚丙酸3-Indolepropionic acid	生长素Auxin	0.00±0.00b	0.55±0.03a	0.00±0.00b	<0.001
tZR	玉米素核苷Trans-Zeatin riboside	细胞分裂素CK	2.27±0.15a	1.60±0.15b	0.40±0.03c	<0.001
BAP9G	6-苄氨基-9-(Α-D-吡喃葡萄糖基)呤 N6-Benzyladenine-9-glucoside	细胞分裂素CK	0.00±0.00b	0.11±0.01a	0.11±0.01a	<0.001
mT9G	3-[[(9-BETA-D-吡喃葡萄糖基-9H-嘌呤-6-基)氨基]甲基]苯酚meta-Topolin-9-glucoside	细胞分裂素CK	0.00±0.00b	0.26±0.04a	0.32±0.05a	0.002
iP9G	异戊烯腺嘌呤-9-葡糖苷N6-Isopentenyl-adenine-9-glucoside	细胞分裂素CK	0.00±0.00c	1.71±0.20b	4.40±0.34a	<0.001
GA19	赤霉素19 Gibberellin A19	赤霉素GA	0.73±0.12a	0.00±0.00b	0.00±0.00b	<0.001
JA	茉莉酸Jasmonic acid	茉莉酸JA	100.61±8.17a	37.79±2.67b	36.24±4.34b	<0.001
OPC-4	氧化戊烯基环戊烷丁酸 3-oxo-2-(2-(Z)-Pentenyl) cyclopentane-1-butyric acid	茉莉酸JA	45.98±3.93a	12.21±0.70b	18.33±1.96b	<0.001
JA-ILE	茉莉酸-异亮氨酸Jasmonoyl-L-isoleucine	茉莉酸JA	8.09±0.71a	4.56±0.05b	0.00±0.00c	<0.001
MeJA	茉莉酸甲酯Methyl jasmonate	茉莉酸JA	2.84±0.08a	0.00±0.00b	0.00±0.00b	<0.001
JA-Val	茉莉酸-缬氨酸N-[(-)-Jasmonoyl]-(L)-valine	茉莉酸JA	0.11±0.004a	0.00±0.00b	0.00±0.00b	<0.001
ABA-GE	脱落酸葡萄糖酯ABA-glucosyl ester	脱落酸ABA	21.61±1.07a	0.00±0b	0.00±0b	<0.001

表3

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序号No	品种（系）Varieties (Lines)	冬性Winterness	选育单位Source
1	GAU30	强冬性Strong winterness	甘肃农业大学Gansu Agricultural University
2	NTS309	强冬性Strong winterness	甘肃农业大学Gansu Agricultural University
3	GAU24	强冬性Strong winterness	甘肃农业大学Gansu Agricultural University
4	16-2444	强冬性Strong winterness	甘肃农业大学Gansu Agricultural University
5	天油14号Tianyou 14	冬性Winterness	天水市农业科学研究所Tianshui Agricultural Science Research Institute
6	天油2238 Tianyou 2238	冬性Winterness	天水市农业科学研究所Tianshui Agricultural Science Research Institute
7	天油2288 Tianyou 2288	冬性Winterness	天水市农业科学研究所Tianshui Agricultural Science Research Institute
8	甘杂9号Ganza 9	弱冬性Weak winterness	西北农林科技大学Northwest Agriculture & Forestry University
9	秦早1号Qinzao 1	弱冬性Weak winterness	陕西省杂交油菜中心Shaanxi Hybrid Rapeseed Center

品种 Varieties	越冬率 Overwintering rate (%)	根颈直径 Collar diameter (mm)	干重 Dry weight (g)	根/冠 Root/crown	生长点高度 The height of growth point (cm)
GAU30	75.00±3.06a	26.38±0.20ab	63.84±6.07ab	0.15±0.02ab	3.17±0.18de
NTS309	74.00±2.31a	27.40±1.49ab	67.41±8.99ab	0.15±0.03ab	3.07±0.25e
GAU24	73.67±2.33a	25.63±1.59b	71.41±2.93ab	0.14±0.03ab	3.27±0.14 cde
16-2444	74.33±2.19a	25.61±1.94b	52.65±6.68b	0.20±0.03a	3.13±0.29de
天油14号Tianyou 14	65.00±3.21b	27.01±0.85ab	80.82±4.53a	0.16±0.04ab	4.06±0.12abc
天油2238 Tianyou 2238	61.67±2.19b	23.20±2.06bc	75.17±10.94ab	0.12±0.01ab	3.96±0.06bcd
天油2288 Tianyou 2288	64.00±2.08b	19.04±0.97c	50.54±8.15b	0.11±0.01b	4.14±0.57ab
甘杂9号Ganza 9	57.00±2.53bc	25.77±0.40b	57.16±4.03ab	0.14±0.01ab	4.75±0.17ab
秦旱1号Qinzao 1	49.67±2.60c	30.65±1.96a	63.80±2.41ab	0.10±0.000b	4.89±0.25a
与越冬率的相关性 The correlation with overwintering rate	1	-0.191(P=0.622)	0.012 (P=0.976)	0.677* (P=0.045)	-0.972** (P<0.001)

甘蓝型冬油菜越冬期矮生长点形态建成及多激素调控机制

Morphogenesis of the Low-Growth Point and Its Multi-Hormonal Regulatory Mechanism During Overwintering in Winter Rapeseed (Brassica napus L.)

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