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作物学报 ›› 2024, Vol. 50 ›› Issue (4): 1015-1029.doi: 10.3724/SP.J.1006.2024.34116

• 耕作栽培·生理生化 • 上一篇    下一篇

甘蓝型油菜苗期响应渍害胁迫的生理调控机制

周香玉(), 徐劲松, 谢伶俐(), 许本波(), 张学昆   

  1. 湿地生态与农业利用教育部工程研究中心 / 长江大学, 湖北荆州 434025
  • 收稿日期:2023-07-06 接受日期:2023-10-23 出版日期:2024-04-12 网络出版日期:2023-11-15
  • 通讯作者: * 谢伶俐, E-mail: linglixie@yangtzeu.edu.cn;许本波, E-mail: benboxu@yangtzeu.edu.cn
  • 作者简介:E-mail: 2844804442@qq.com
  • 基金资助:
    农业生物育种重大项目(2023ZD04042);农业农村部项目(15214011);湖北省农业厅项目(鄂农油[2022]7号)

Physiological mechanisms in response to waterlogging during seedling stage of Brassica napus L.

ZHOU Xiang-Yu(), XU Jin-Song, XIE Ling-Li(), XU Ben-Bo(), ZHANG Xue-Kun   

  1. Engineering Research Center of Ecology and Agricultural Use of Wetland, Ministry of Education / Yangtze University, Jingzhou 434025, Hubei, China
  • Received:2023-07-06 Accepted:2023-10-23 Published:2024-04-12 Published online:2023-11-15
  • Contact: * E-mail: linglixie@yangtzeu.edu.cn; E-mail: benboxu@yangtzeu.edu.cn
  • Supported by:
    Major Projects of Agricultural Biology Breeding of China(2023ZD04042);Entrusted Program of Ministry of Agriculture and Rural Affairs of China(15214011);Hubei Provincial Department of Agriculture Project (Hubei Province Nongyou [2022] 7)

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摘要：

长江流域是中国油菜主产区, 该区域常年湿润多雨, 且产区实行油菜-水稻轮作制度, 导致渍害频发。为明确甘蓝型油菜(Brassica napus L.)对苗期渍害的响应机制, 本研究采用盆栽试验, 以强耐渍品系YZ12、中等耐渍品系YZ45和不耐渍品系YZ59为试验材料, 研究苗期淹水对油菜表型性状、生理特性、光合作用、相关基因相对转录水平等的影响, 同时分析了外源激素抑制剂对油菜渍害胁迫的影响。结果表明, 淹水胁迫严重抑制油菜生长, 根系活力可作为衡量淹水胁迫对油菜生长影响的指示指标。根细胞超微结构观察发现, 淹水胁迫导致油菜根系细胞发生质壁分离及细胞器破碎解体, 强、中耐渍油菜的细胞器受损程度较小, 能够在淹水胁迫中维持较为正常的细胞形态; 淹水胁迫下根部细胞骨架相关基因Bnamicrotubule1.A3、Bnatubulin-α2.C3、Bnatubulin-β7.C6、Bnalamin-like.A2相对转录水平显著下调至对照水平(CK)的0.2~0.5倍; 无氧呼吸相关基因BnaPDC.C9、BnaLDH.A1、BnaADH.A7表达量显著升高, 为CK的3~6倍, 且在中、强耐渍油菜中诱导表达水平更高。过氧化物酶(peroxidase, POD)、超氧化物歧化酶(superoxide dismutase, SOD)活性随淹水时间延长呈先升后降趋势, 过氧化氢酶(catalase, CAT)活性和丙二醛(malondialdehyde, MDA)含量呈升高趋势, 其中强耐渍品系抗氧化酶活性相对较高, 而MDA增幅较小。淹水胁迫严重影响油菜叶片光合效率及叶绿素含量, 导致油菜叶绿素含量、光合速率、气孔导度和蒸腾速率显著下降, 胞间CO2浓度显著升高, 且不耐渍品系变化幅度相对较大。淹水胁迫导致油菜乙烯(ethylene, ET)和脱落酸(abscisic acid, ABA)含量显著升高, 其中强耐渍品系ET含量较高, 不耐渍品系ABA含量较高; 强耐渍品系的ET信号相关基因BnaACO1.C8、BnaERF73.C6相对转录水平显著上调, 不耐渍品系ABA合成相关基因BnaZEP.A7相对转录水平上调。外源喷施激素抑制剂可改善淹水胁迫对油菜的伤害, 但不同外源激素抑制剂效果差异明显。综上, 不同耐渍性甘蓝型油菜苗期对淹水胁迫响应在表型、生理代谢、光合、激素和基因转录水平存在差异。甘蓝型油菜通过调控细胞骨架、无氧呼吸、激素代谢相关基因的相对转录水平, 引起植株内抗氧化酶活性、激素水平、光合效率、根部超微结构及根系活力改变, 进而响应淹水胁迫。

关键词: 甘蓝型油菜, 淹水胁迫, 根系超微结构, 光合特性, 抗氧化酶活性, 激素水平, 转录调控

Abstract:

The Yangtze River basin is the main producing area of rapeseeds in China, which is wet and rainy all the year round, and the rapeseed-rice rotation system is implemented in the producing area, resulting in frequent waterlogging. To explore the effects of waterlogging at seedling stage on phenotypic traits, physiological characteristics, photosynthesis, relative gene transcriptional levels, and the regulation of exogenous hormone inhibitors on rapeseed damage under waterlogging, a pot experiment was conducted, and the strong waterlogging tolerant line YZ12, medium waterlogging tolerant line YZ45, and weak waterlogging tolerant line YZ59 were used as the experimental materials. The results indicated that flooding stress severely inhibited the growth of rapeseed, and root activity could be used as an indicator to measure the impact of flooding stress on rapeseed growth. The observation of root cell ultrastructure showed that flooding stress led to plasmolysis and organelle fragmentation of rape root cells. The organelle of strong and medium waterlogging resistant rape was less damaged, and it could maintain a more normal cell morphology under flooding stress. The relative transcriptional levels of cytoskeletal genes Bnamicrotubule1.A3, Bnatubulin-α2.C3, Bnatubulin-β7.C6, and Bnalamin-like.A2 in rape roots were significantly decreased under flooding stress, which were 0.2-0.5 times that of the control (CK) samples. The relative expressional levels of BnaPDH.C9, BnaLDH.A1, and BnaADH.A7 associated to anaerobic respiration were significantly increased, which was 3-6 times higher than that of CK, and higher expression levels were observed in medium and strong waterlogging tolerant rapeseed seedlings than in weak waterlogging tolerant line YZ59. During waterlogging, the activities of POD and SOD increased first and then decreased, while the activity of CAT and the content of MDA increased. Among them, the enzyme activities of YZ12 line such as POD, SOD, CAT were relatively high, and the increase of MDA was small. The photosynthetic efficiency and chlorophyll content of rapeseed leaves were seriously affected by flooding stress. The chlorophyll content, photosynthetic rate, stomatal conductance and transpiration rate of rapeseed decreased significantly, and the intercellular CO2 concentration increased significantly, and the change range of the weak waterlogging tolerant line YZ59 was larger than that of the other two lines. Under flooding stress, ET and ABA contents of rapeseed increased significantly. Among the three lines, YZ12 had higher ET content, while YZ59 had higher ABA content. The relative transcriptional levels of ET related genes BnaACO1.C8, BnaERF73.C6 were significantly up-regulated in the strong waterlogging tolerant line YZ12, while the relative transcriptional level of ABA-related gene BnaZEP.A7 was up-regulated in the weak waterlogging tolerant line YZ59. Exogenous application of hormone inhibitors could improve the damage of flooding stress to rapeseed, but the effects of different exogenous hormone inhibitors varied significantly. In conclusion, there were differences in physiological metabolism, photosynthesis, hormone, and gene transcriptional levels in response to flooding stress at seedling stage in B. napus with different waterlogging tolerance. B. napus responsed to flooding stress by regulating the relative transcription levels of genes related to cytoskeleton, anaerobic respiration, hormone metabolism, causing changes in antioxidant enzyme activity, hormone levels, photosynthetic efficiency, root ultrastructure and root vitality.

Key words: Brassica napus L., flooding stress, root ultrastructure, photosynthetic characteristics, antioxidant enzyme activity, hormone levels, transcriptional regulation

表1

qRT-PCR所用引物序列"

基因编号
Gene ID		 基因名称
Gene name		 正向引物
Forward primer (5°-3°)		 反向引物
Reverse primer (5°-3°)
XM_013825609.2 Bnatubulin-α2.C3 AACCATCAAGACCAAGCGCAC CCTCCTCATAATCCTTCTCCAG
XM_013846861.2 Bnatubulin-β7.C6 CGTCCAGAACAAGAACTCCTC GCTCGCTGACTCTCCTGAAC
XM_013847632.2 Bnalaminlike.A2 GCATCTCCGACCATCCTATCCG AAACAACCACCCCTTCCGTC
XM_013786846.2 Bnamicrotubule1.A3 GCATACGATGGTGTTCCTCTG CTCTACGTGTGGCTGTTCTTG
XM_013852099.3 BnaPDC.C9 TGCTCAAGCCATACCTAACAAACG CGCAGTCCAGCATTTACCTTCTC
XM_013794611.3 BnaADH.A7 GGATTGCTGGTGCTGGTAGGA GCGATGACTTGCTGAACTGGC
XM_013866109.3 BnaLDH.A1 TCCTCCGAGACCAGCGTAAGAT TTAGTCACAGCCACCACACCG
GU550519.1 BnaRGA.A9 ATCACTCTCTCTCCGACACGCTC GGACCACCTTCTCTCAACGCAA
XM_013788321.3 BnaGA3.A6 GGATAAGAAGCGTTGGGAGAAGC TCAACATTCTCTTCCTCACCGTCT
XM_013815976.3 BnaGA2ox2.C3 TGAAAGATGGAAGTTGGGTCGC GCAATCTTCTGGCTCAATGGG
NM_001315888.1 BnaZEP.A7 GCATTGTCGGAAGTGAACCAGA TGAGATAGGTCCCGTGTTCGC
XM_013788810.3 BnaNCED3.C1 ACCAGATACGCTTACCTCGCTT TGTAACCGTCGTCTTCGCCT
XM_013887010.1 BnaAAO.A4 GATTTAGGACAGGTGGAAGGAGC CACAATGAACCGAAGCCGC
XM_013804747.3 BnaACO1.C8 CGCCATCAAAGAACAACACCATT CTGGAGCTGGAGATATTACGGCA
XM_013893836.3 BnaEIN3.A5 CCAACGGTCCAGAATCATCAAGA GTTGTTGTTGTTGTTGGTGTGCG
XM_013845804.2 BnaERF73.C6 CAACTTCCCTAACGATTCTCCAGC CCGCATTGTTATTCTCCTCCCAC
NM_001316010.1 BnaACT.C2 CTGGAATTGCTGACCGTATGAG ATCTGTTGGAAAGTGCTGAGGG

图1

淹水胁迫6 d不同耐渍性甘蓝型油菜形态 CK: 正常供水; T: 淹水处理6 d。"

表2

淹水6 d时3个油菜品系形态指标及根系活力"

品系
Line		 处理
Treatment		 株高
Plant height
(cm)		 叶长
Leaf length
(cm)		 叶宽
Leaf width
(cm)		 地上部鲜重
Fresh weight of shoot (g)		 根长
Root length
(cm)		 根系活力
Root activity
(mg g-1 h-1)
YZ12 CK 13.63±1.42 a 4.40±0.39 a 3.87±0.47 a 1.07±0.16 a 7.93±1.17 a 453.59±12.71 a
T 10.17±0.89 bc 2.40±0.38 c 2.07±0.16 c 0.70±0.02 bc 5.52±1.67 bc 385.07±23.59 b
YZ45 CK 11.35±1.70 b 3.90±0.86 ab 3.47±0.19 b 0.91±0.27 ab 6.85±0.96 ab 442.02±14.39 a
T 8.80±0.63 cd 2.25±0.21 c 1.98±0.27 c 0.55±0.06 c 3.97±1.14 cd 364.69±8.26 bc
YZ59 CK 10.98±1.39 b 3.57±0.31 b 3.53±0.40 ab 0.90±36.00 ab 8.08±1.78 a 462.23±35.66 a
T 8.27±1.01 d 1.90±0.12 c 1.72±0.12 c 0.26±0.18 c 3.12±1.17 c 325.93±26.33 c

表3

淹水6 d时3个油菜品系各指标的灰色关联分析"

指标
Index		 根系活力
Root activity		 叶宽
Leaf width		 根长
Root length		 叶长
Leaf length		 株高
Plant height
关联度 Correlation degree 0.6211 0.6001 0.5938 0.5798 0.5465
排序 Order 1 2 3 4 5

图2

不同耐渍性甘蓝型油菜根细胞超微结构 CK: 正常供水; T: 淹水处理6 d; A~C: 正常供水状态下不同耐渍性油菜根细胞超微结构; D~F: 淹水胁迫下不同耐渍性油菜根细胞超微结构。"

图3

根中细胞骨架相关基因转录水平分析 CK: 正常供水; T: 淹水处理48 h。不同小写字母表示处理间在0.05概率水平差异显著。"

图4

根中无氧呼吸相关基因转录水平分析 CK: 正常供水; T: 淹水处理48 h。不同小写字母表示处理间在0.05概率水平差异显著。"

图5

淹水胁迫下不同耐渍性甘蓝型油菜内源激素含量的变化"

图6

根中激素代谢相关基因转录水平分析 CK: 正常供水; T: 淹水处理48 h。不同小写字母表示处理间在0.05概率水平差异显著。"

图7

淹水胁迫对甘蓝型油菜抗氧化酶活性及MDA含量的影响 不同小写字母表示处理间在0.05概率水平差异显著。"

图8

淹水胁迫对甘蓝型油菜光合性状及叶绿素总量的影响 不同小写字母表示处理间在0.05概率水平差异显著。"

图9

甘蓝型油菜根系活力与生理指标相关性分析 图中数据为Pearson r。X1: 根系活力; X2: POD活性; X3: CAT活性; X4: SOD活性; X5: MDA含量; X6: 光合速率; X7: 气孔导度; X8: 蒸腾速率; X9: 胞间CO2浓度; X10: 叶绿素含量; X11: GA含量; X12: ABA含量; X13: ET含量。"

图10

外源生长调节剂对淹水胁迫下甘蓝型油菜表型的影响 CK: 正常供水; T1: 淹水处理+喷施蒸馏水; T2: 淹水处理+喷施GA抑制剂UNICONAZOLE; T3: 淹水处理+喷施ET抑制剂STS; T4: 淹水处理+喷施ABA抑制剂NDGA。"

图11

淹水胁迫对甘蓝型油菜苗期生长的影响及响应机制 A: 激素调控根系生长信号通路图[12,50??-53], 实线表示直接相互作用, 虚线表示间接相互作用。箭头表示刺激作用, ⊥表示抑制作用; B: 甘蓝型油菜苗期响应渍害胁迫的生理调控机制。"

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