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作物学报 ›› 2021, Vol. 47 ›› Issue (10): 2001-2011.doi: 10.3724/SP.J.1006.2021.04238

• 耕作栽培·生理生化 • 上一篇    下一篇

盐胁迫下外源2,4-表油菜素内酯对颠茄氮代谢及TAs代谢的影响

辛正琦1(), 代欢欢2, 辛余凤1, 何潇1, 谢海艳1, 吴能表1,*()   

  1. 1西南大学生命科学学院 / 三峡库区生态环境教育部重点实验室, 重庆 400715
    2甘肃庆阳市第五中学, 甘肃庆阳 745000
  • 收稿日期:2020-11-01 接受日期:2021-01-13 出版日期:2021-10-12 网络出版日期:2021-02-25
  • 通讯作者: 吴能表
  • 作者简介:E-mail: 490992699@qq.com
  • 基金资助:
    国家自然科学基金项目(30500041)

Effects of exogenous 2,4-Epibrassinolide on nitrogen metabolism and TAs metabolism of Atropa belladonna L. under NaCl stress

XIN Zheng-Qi1(), DAI Huan-Huan2, XIN Yu-Feng1, HE Xiao1, XIE Hai-Yan1, WU Neng-Biao1,*()   

  1. 1School of Life Science, Southwest University / Key Laboratory of Eco-environments in Three Gorges Region, Ministry of Education, Chongqing 400715, China
    2The Fifth Middle School of Qingyang, Qingyang 745000, Gansu, China
  • Received:2020-11-01 Accepted:2021-01-13 Published:2021-10-12 Published online:2021-02-25
  • Contact: WU Neng-Biao
  • Supported by:
    National Natural Science Foundation of China(30500041)

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摘要：

以颠茄(Atropa belladonna L.)幼苗为材料, 采用盆栽试验, 使用外源2,4-表油菜素内酯(2,4-Epibrassinolide, EBR)处理幼苗, 研究在100 mmol L-1 NaCl胁迫下不同浓度(0.05、0.1、0.2、0.4 mg L-1)外源EBR在不同处理时间(5、10、15、20 d)内对颠茄氮代谢、次生代谢产物含量以及托品烷类生物碱(tropane alkaloids, TAs)合成途径中前体物质含量、关键酶基因表达量的影响, 旨在明确外源EBR调控颠茄耐盐性的生理机制。盐胁迫下颠茄氮代谢受到抑制, 而施加外源EBR能够有效增强颠茄的氮代谢能力, 硝态氮含量显著升高, 铵态氮含量降低, 游离氨基酸、可溶性蛋白含量以及氮代谢关键酶活性均有不同程度的上升。盐胁迫不利于生物碱的合成和积累, 显著降低了TAs途径中前体物质的合成和关键酶基因的表达。外施0.1 mg L-1EBR能够有效提高鸟氨酸、精氨酸、多胺含量以及腐胺合成关键酶活性, 并通过上调TAs途径中关键酶基因TR I、H6H的表达量来提高莨菪碱和东莨菪碱的含量。表明适宜浓度的外源EBR能够有效缓解盐胁迫对颠茄生理代谢的破坏, 通过提高氮代谢能力、促进TAs的产生和积累, 提高颠茄幼苗的耐盐性。

关键词: 颠茄, 盐胁迫, 氮代谢, 次生代谢

Abstract:

To explore the physiological mechanism of exogenous 2,4-Epibrassinolide (EBR) regulating NaCl tolerance of A. belladonna, potted A. belladonna seedlings was used as the experimental materials, and exogenous EBR was applied to A. belladonna seedlings with the different concentrations (0.05, 0.1, 0.2, and 0.4 mg L-1) of exogenous EBR and different treatment times (5, 10, 15, and 20 days) on the nitrogen metabolism, the contents of the secondary metabolites and precursor substances in the synthesis pathway of TAs, and the relative expression levels of key enzyme genes. NaCl stress caused inhibitory effect on nitrogen metabolism in A. belladonna, however the content of nitrate nitrogen increased significantly, the content of ammonium nitrogen decreased, the content of free amino acids, soluble protein, and the activity of key enzymes of nitrogen metabolism increased to some extent under exogenous EBR treatment, which indicating the exogenous EBR could effectively enhance the nitrogen metabolism capacity. NaCl stress was not conducive to the synthesis and accumulation of alkaloids. The synthesis of precursor substances and the relative expression levels of key enzyme genes in TAs pathway significantly were reduced under NaCl stress. The contents of ornithine, arginine, polyamine, and the activities of key enzymes in putrescine synthesis were increased with the exogenous EBR of 0.1 mg L-1. Moreover, exogenous EBR could effectively enhance the contents of hyoscyamine and scopolamine by increasing the relative expression levels of key enzyme genes TR I and H6H in TAs pathway. In conclusion, the appropriate concentration of exogenous EBR could effectively relieve the damage of NaCl stress to the physiological metabolism in A. belladonna, and the exogenous EBR could improve NaCl tolerance of belladonna seedlings by increasing nitrogen metabolism and promoting the production and accumulation of TAs.

Key words: Atropa belladonna L., NaCl stress, nitrogen metabolism, secondary metabolism

表1

颠茄幼苗的不同处理组合"

处理组
Group		 NaCl盐浓度
NaCl concentration (mmol L-1)		 EBR浓度
EBR concentration
(mg L-1)
CK 0 0
T0 100 0
B1 100 0.05
B2 100 0.1
B3 100 0.2
B4 100 0.4

表2

荧光定量PCR检测所用引物"

引物名称
Primer name		 引物序列
Primer sequence (5'-3')
F-qPGK TCGCTCTTGGAGAAGGTTGAC
R-qPGK CTTGTCCGCAATCACTACATCAG
F-AbTR Ⅰ TTCTTTGCTTCCCTGCTGCTTC
R-AbTR Ⅰ CAGGCCAACCTTAGTATCACACAG
F-AbH6H TTCCACTTGAGCAGAAAGCAAAGC
R-AbH6H CCTCATGGTCAACTTCCTCACTTCC
F-AbPMT CCTACTTACCCTACTGGTGTTATC
R-AbPMT GCGAAAGATGGCAAAATAAAAGC
F-AbPPAR GATTGGAGTTGTTGGATTGG
R-AbPPAR CCTTGAAGTGTAAGAGATGGA

图1

盐胁迫20 d时0.1 mg L-1外源EBR处理后颠茄幼苗的生长情况 处理同表1。标尺为10 cm。"

表3

不同浓度外源EBR对盐胁迫下颠茄叶片含氮化合物含量的影响"

指标
Indicator		 处理组
Group		 处理时间Treatment days
5 d 10 d 15 d 20 d
硝态氮
Nitrate nitrogen
(μg g-1 FW)		 CK 235.270±19.155 a 219.531±17.631 a 228.121±17.588 a 223.681±10.459 a
T0 172.379±10.148 c 165.078±13.045 d 144.347±15.771 d 126.918±15.774 d
B1 196.475±15.896 bc 184.987±15.751 cd 171.979±18.628 cd 155.752±12.174 c
B2 221.515±18.481 ab 212.715±25.806 ab 192.219±10.863 bc 192.379±16.613 b
B3 238.206±17.135 a 206.361±16.391 ab 209.203±15.735 ab 180.566±18.503 bc
B4 209.142±23.651 ab 193.902±15.395 bc 184.267±11.701 bc 177.438±15.189 bc
铵态氮
Ammonium nitrogen
(μg g-1 FW)		 CK 86.897±4.105 a 76.944±2.499 c 67.929±2.828 d 71.087±4.672 d
T0 72.957±2.013 c 91.748±1.776 a 106.025±1.289 a 124.603±5.316 a
B1 81.997±1.921 ab 83.524±3.315 b 90.275±1.066 b 113.194±1.749 ab
B2 78.304±3.566 bc 74.326±2.223 cd 71.299±3.039 d 82.622±7.815 cd
B3 73.251±0.933 c 69.352±5.986 d 81.022±3.893 c 93.441±3.973 bc
B4 77.815±2.184 bc 79.089±3.618 bc 86.931±5.621 bc 98.837±5.552 bc
游离氨基酸
Free amino acids
(μg g-1 FW)		 CK 821.721±30.434 a 784.398±26.959 a 797.848±43.402 a 758.979±36.011 a
T0 691.934±11.070 c 651.923±5.596 d 606.195±35.981 d 586.021±28.179 c
B1 740.688±15.033 bc 685.546±18.517 cd 647.551±15.952 cd 663.018±11.939 b
B2 782.717±20.377 ab 716.479±14.387 bc 732.619±16.309 ab 708.746±20.148 ab
B3 754.810±13.649 b 741.024±11.110 ab 744.723±17.181 ab 680.879±17.181 b
B4 728.920±8.578 bc 704.375±12.221 bc 692.271±6.623 bc 654.612±9.047 bc
可溶性蛋白
Soluble proteins
(mg g-1 FW)		 CK 2.926±0.072 c 2.721±0.085 bc 2.625±0.121 a 2.501±0.058 a
T0 2.839±0.241 bc 2.542±0.109 c 1.816±0.094 c 1.667±0.050 d
B1 3.021±0.094 ab 2.741±0.095 bc 1.979±0.129 bc 1.927±0.104 cd
B2 3.361±0.110 a 3.002±0.051 ab 2.374±0.234 ab 2.308±0.144 ab
B3 3.122±0.071 ab 3.106±0.118 a 2.557±0.146 a 2.149±0.119 bc
B4 2.895±0.104 bc 3.038±0.142 ab 2.056±0.122 bc 2.095±0.126 bc

表4

不同浓度外源EBR对盐胁迫下颠茄叶片硝酸还原酶, 谷氨酰胺合成酶和谷氨酸脱氢酶活性的影响"

指标
Indicator		 处理组
Group		 处理时间Treatment days
5 d 10 d 15 d 20 d
硝酸还原酶
NR
(U g-1 FW h-1)		 CK 1.445±0.028 ab 1.396±0.065 ab 1.462±0.027 a 1.429±0.037 a
T0 1.233±0.063 d 1.065±0.074 d 0.932±0.104 d 0.798±0.076 d
B1 1.392±0.041 cd 1.174±0.045 cd 0.974±0.070 cd 0.874±0.053 d
B2 1.394±0.017 bc 1.455±0.073 a 1.251±0.926 b 1.185±0.063 b
B3 1.566±0.043 a 1.396±0.026 ab 1.149±0.060 bc 1.049±0.052 bc
B4 1.511±0.055 ab 1.257±0.028 bc 1.106±0.026 cd 0.939±0.074 cd
谷氨酰胺合成酶
GS
(OD540 mg-1 protein FW h-1)		 CK 3.676±0.126 a 3.482±0.172 a 3.304±0.168 a 3.392±0.067 a
T0 2.992±0.154 c 2.733±0.164 c 2.467±0.143 d 2.303±0.133 d
B1 3.148±0.048 bc 2.867±0.102 bc 2.698±0.081 cd 2.465±0.085 cd
B2 3.570±0.140 a 3.187±0.173 ab 3.225±0.102 ab 3.044±0.036 b
B3 3.512±0.067 ab 3.437±0.042 a 3.025±0.108 bc 2.698±0.124 c
B4 3.429±0.102 ab 3.258±0.080 ab 2.889±0.098 bc 2.527±0.113 cd
谷氨酸脱氢酶
GDH
(U mg-1 FW min-1)		 CK 0.252±0.026 c 0.271±0.016 bc 0.772±0.016 b 0.283±0.033 ab
T0 0.293±0.019 bc 0.202±0.008 d 0.154±0.025 d 0.184±0.027 c
B1 0.271±0.024 bc 0.241±0.027 cd 0.198±0.017 cd 0.230±0.019 bc
B2 0.356±0.025 a 0.305±0.022 ab 0.283±0.026 b 0.305±0.024 ab
B3 0.315±0.022 ab 0.346±0.026 a 0.359±0.034 a 0.362±0.016 a
B4 0.236±0.163 c 0.242±0.031 cd 0.258±0.037 bc 0.248±0.025 bc

图2

不同浓度外源EBR对盐胁迫下颠茄叶片莨菪碱(A)和东莨菪碱(B)含量的影响 不同小写字母表示处理间在0.05水平差异显著。处理同表1。"

图3

不同浓度外源EBR对盐胁迫下20 d时颠茄叶片鸟氨酸和精氨酸含量的影响 不同小写字母表示处理间在0.05水平差异显著。处理同表1。"

图4

不同浓度外源EBR对盐胁迫下20 d时颠茄叶片多胺含量(A)、多胺总量(B)的影响 不同小写字母表示处理间在0.05水平差异显著。处理同表1。"

图5

不同浓度外源EBR对盐胁迫下20 d时颠茄叶片多胺合成关键酶活性的影响 不同小写字母表示处理间在0.05水平差异显著。处理同表1。"

图6

不同浓度外源EBR对盐胁迫下颠茄根和叶中TR I (A)、PMT (B)、H6H (C)、PPAR (D)相对基因表达量的影响 不同小写字母表示处理间在0.05水平差异显著。处理同表1。"

图7

盐胁迫20 d 时0.1 mg L-1外源EBR处理后颠茄氮代谢及TAs代谢途径网络变化 红色代表显著上升; 绿色代表显著降低; 白色代表无明显变化; FAA: 游离氨基酸; SP: 可溶性蛋白; Orn: 鸟氨酸; Arg: 精氨酸; Hyo: 莨菪碱; Sco: 东莨菪碱。"

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