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作物学报 ›› 2018, Vol. 44 ›› Issue (10): 1527-1538.doi: 10.3724/SP.J.1006.2018.01527

• 耕作栽培·生理生化 • 上一篇    下一篇

UV-B胁迫下Ca 2+对颠茄生理特性与次生代谢产物的调控研究

卢克欢1,刘兴1,杨怡1,廖志华2,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西南大学生命科学学院, 重庆 400715
  • 收稿日期:2018-01-15 接受日期:2018-07-20 出版日期:2018-10-10 网络出版日期:2018-08-01
  • 通讯作者: 吴能表
  • 基金资助:
    本研究由国家自然科学基金项目(30500041);重庆市科技攻关计划项目(cstc2012gg-yyjs80013)

Effect of Exogenous Ca 2+ on Physiological Characteristics and Secondary Metabolites accumulation of Atropa belladonna L. Seedlings under UV-B Stress

Ke-Huan LU1,Xing LIU1,Yi YANG1,Zhi-Hua LIAO2,Neng-Biao WU1,*()   

  1. 1 Key Laboratory of Eco-environments in Three Gorges Reservoir Region, Ministry of Education / School of Life Science, Southwest University, Chongqing 400715, China
    2 School of Life Science, Southwest University, Chongqing 400715, China
  • Received:2018-01-15 Accepted:2018-07-20 Published:2018-10-10 Published online:2018-08-01
  • Contact: Neng-Biao WU
  • Supported by:
    This study was supported by the National Natural Science Foundation of China(30500041);the Science Technology Breakthrough Plan Project of Chongqing(cstc2012gg-yyjs80013)

摘要:

有害中波紫外线(Ultraviolet B, UV-B; 280~320 nm)辐射影响植物的生长和发育, Ca 2+是植物生长发育所必须的大量元素之一, 外源Ca 2+不但可以提高植物抗胁迫能力, 还对次生代谢具有调控作用。以颠茄(Atropa belladonna L.)实生苗为材料, 在UV-B辐射(10 μW cm -2)背景下, 研究不同浓度外源Ca 2+、不同处理时间对颠茄生理特性、氮代谢、次生代谢产物含量以及托品烷类生物碱代谢途径中3个关键酶基因表达量的影响。结果表明, 随着UV-B辐射时间的延长(4 ~12 d), 对颠茄的光合作用、氮代谢以及生物碱的积累产生抑制作用, 加剧了膜脂氧化程度; 经外源Ca 2+处理, 叶片初始荧光(Fo)、丙二醛(malondialdehyde, MDA)含量呈下降趋势, 叶片最大光化学效率(Fv/Fm)、光合色素(总叶绿素、类胡萝卜素)含量、抗氧化酶(SOD、POD、CAT)活性均呈上升趋势, 说明Ca 2+有利于缓解UV-B辐射的抑制, 加强对UV-B胁迫的抗性; 在Ca 2+处理下, 叶片中硝态氮含量显著降低, 游离氨基酸、可溶性蛋白含量和氮代谢关键酶(NR、GS、GDH)的活性显著提高, 叶片中莨菪碱含量和东莨菪碱含量显著提高; qRT-PCR分析显示, 在外源Ca 2+的诱导下, 托品烷类生物碱合成途径中3个关键酶基因(PMTTR IH6H)表达量均有不同程度上调趋势。本研究结果可为田间种植提供理论参考。

关键词: UV-B胁迫, 颠茄, 生理特性, 氮代谢, 托品烷类生物碱

Abstract:

Ultraviolet B (UV-B) radiation does harm to plant, for which growth and development. Ca 2+is one of the most necessary elements. Exogenous Ca 2+ can enhance plant abilities to resist stress and regulate secondary metabolism. In this research, we used Atropa belladonna L. seedlings to investigate the effects of different concentrations of exogenous Ca 2+ and different treatment days on the physiological characteristics, nitrogen metabolism, secondary metabolites content and relative genes expression levels of three key enzymes in secondary metabolism under the background of UV-B radiation. The UV-B radiation not only caused inhibitory effect on the Atropa belladonna L. photosynthesis, nitrogen metabolism, alkaloid content, but also increased peroxidation of membrane lipid. The Fo, MDA content gradually decreased, but the Fv/Fm, photosynthetic pigment content, antioxidant enzyme activity gradually increased under Ca 2+ treatment, showing that Ca 2+ is propitious to relieve inhibitory effect on photosynthesis and enhance resistance to UV-B stress. Moreover, Ca 2+ reduced the content of nitrate nitrogen significantly, promoted the contents of free amino acids, soluble proteins, atropine alkaloids, and raised the activities of key enzymes in nitrogen metabolism. Using real-time fluorescence quantitative PCR, we found that exogenous Ca 2+ increased the relative gene expression levels of three key enzymes in secondary metabolism in different degrees. This research could provide a theoretical references for plantation in the field.

Key words: UV-B stress, Atropa belladonna L., physiological characteristics, nitrogen metabolism, tropane alkaloids

表1

颠茄幼苗的处理组合"

处理组
Group
CaCl2 浓度
CaCl2 concentration (mmol L-1)
UV-B辐射强度
UV-B radiation intensity (μW cm-2)
处理时间
Treatment time (d)
T0 0 0 4, 8, 12
T1 0 10 4, 8, 12
T2 2 10 4, 8, 12
T3 4 10 4, 8, 12
T4 6 10 4, 8, 12
T5 8 10 4, 8, 12

表2

荧光定量PCR所需引物"

基因
Gene
登录号
Accession number
上游引物
Forward primer (5'-3')
下游引物
Reverse primer (5'-3')
退火温度
Annealing temperature (°C)
PGK JX154676 TCGCTCTTGGAGAAGGTTGAC CTTGTCCGCAATCACTACATCAG 59.5
PMT AB018570 CCTACTTACCCTACTGGTGTTATC GCGAAAGATGGCAAAATAAAAGC 56.4
H6H JN542537 TTCCACTTGAGCAGAAAGCAAAGC CCTCATGGTCAACTTCCTCACTTCC 56.4
TR I JX155757 TTCTTTGCTTCCCTGCTGCTTC CAGGCCAACCTTAGTATCACACAG 61.4

图1

UV-B胁迫下不同浓度Ca2+对颠茄初始荧光(A)和最大光化学效率(B)的影响 不同小写字母表示相同处理时间不同浓度Ca2+处理间差异显著(P<0.05)。T1至T5分别代表UV-B胁迫下不同浓度Ca2+处理组, 分别为0 mmol L-1、2 mmol L-1、4 mmol L-1、6 mmol L-1和8 mmol L-1。T0表示未进行任何处理, 即空白对照组。"

图2

UV-B胁迫下不同浓度Ca2+对颠茄叶片中总叶绿素含量(A)与类胡萝卜素含量(B)的影响 不同小写字母表示相同处理时间不同浓度Ca2+处理间差异显著(P < 0.05)。T1至T5分别代表UV-B胁迫下不同浓度Ca2+处理组, 分别为0、2、4、6和8 mmol L-1。T0表示未进行任何处理, 即空白对照组。"

图3

UV-B胁迫下不同浓度Ca2+对颠茄抗氧化酶活性(A, B, C)、丙二醛含量(D)的影响 不同小写字母表示相同处理时间不同浓度Ca2+处理间差异显著(P < 0.05)。T1至T5分别代表UV-B胁迫下不同浓度Ca2+处理组, 分别为0、2、4、6和8。T0表示未进行任何处理, 即空白对照组。"

表3

UV-B胁迫下不同浓度Ca2+对颠茄硝态氮、游离氨基酸和可溶性蛋白含量的影响"

指标
Indicator
处理组
Group
处理时间 Treatment time
4 d 8 d 12 d
硝态氮
Nitrate nitrogen
(μg g-1 FW)
T0 539.543±19.299 b 545.976±29.480 a 526.677±29.480 a
T1 693.935±38.598 d 874.059±58.959 c 1112.080±40.174 d
T2 610.306±40.174 c 803.296±29.480 c 944.822±57.897 c
T3 462.347±19.299 a 726.100±48.568 b 874.059±62.038 c
T4 507.378±48.568 ab 597.440±38.598 a 783.997±29.480 b
T5 494.512±29.480 ab 681.069±44.569 b 861.193±48.568 bc
游离氨基酸
Free amino acids
(mg g-1 FW)
T0 352.269±32.459 e 329.930±15.145 d 332.723±14.323 d
T1 225.763±38.646 b 177.929±25.124 a 134.465±19.857 a
T2 287.559±33.889 cd 185.213±25.860 a 161.661±18.992 ab
T3 313.176±25.550 de 255.872±30.828 bc 241.789±24.322 c
T4 251.137±33.889 bc 286.224±22.980 cd 264.856±17.015 c
T5 166.153±29.940 a 221.635±35.763 ab 194.076±16.096 b
指标
Indicator
处理组
Group
处理时间 Treatment time
4 d 8 d 12 d
可溶性蛋白
Soluble proteins
(mg g-1 FW)
T0 5.879±0.309 a 5.681±0.364 ab 5.570±0.398 c
T1 6.458±0.401 a 4.953±0.548 a 3.725±0.430 a
T2 7.901±0.653 bc 5.736±0.432 ab 5.264±0.649 bc
T3 9.087±0.399 c 7.415±0.572 c 6.953±0.391 d
T4 6.859±0.491 ab 7.496±0.749 c 4.358±1.058 ab
T5 5.861±1.526 a 6.535±1.007 bc 3.887±0.456 a

表4

UV-B胁迫下不同浓度Ca2+对颠茄硝酸还原酶、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 time
4 d 8 d 12 d
硝酸还原酶 NR
(U g-1 FW h-1)
T0 1.290±0.048 c 1.282±0.117 c 1.333±0.052 d
T1 1.037±0.022 a 0.813±0.053 a 0.557±0.039 a
T2 1.035±0.061 a 0.939±0.108 a 0.664±0.040 b
T3 1.121±0.009 b 1.083±0.030 b 0.724±0.047 b
T4 1.290±0.018 c 1.213±0.035 bc 0.902±0.043 c
T5 1.135±0.013 b 1.121±0.048 b 0.753±0.080 b
谷氨酰胺合成酶 GS
(OD540 mg-1 FW h-1)
T0 8.349±0.197 c 7.885±0.259 d 7.696±0.670 d
T1 6.555±0.235 a 4.989±0.767 a 3.952±0.323 a
T2 6.812±0.297 a 5.666±0.155 b 4.323±0.214 ab
T3 7.494±0.285 b 6.442±0.138 c 4.883±0.226 bc
T4 8.037±0.231 c 7.336±0.189 d 5.454±0.201 c
T5 6.870±0.268 a 6.104±0.105 bc 4.592±0.219 ab
谷氨酸脱氢酶 GDH
(U mg-1 FW min-1)
T0 5.659±0.402 a 5.756±0.658 b 4.968±0.482 b
T1 6.109±0.589 ab 4.244±0.579 a 3.344±0.486 a
T2 6.367±0.841 ab 5.338±0.486 ab 4.887±0.402 b
T3 7.138±0.880 b 5.916±0.990 b 6.881±0.620 c
T4 6.045±0.780 ab 8.039±0.678 c 6.431±0.403 c
T5 5.016±0.696 a 7.395±0.486 c 5.338±0.589 b

表5

UV-B胁迫下不同浓度Ca2+对颠茄叶片莨菪碱和东莨菪碱含量的影响"

指标
Indicator
处理组
Group
处理时间 Treatment time
4 d 8 d 12 d
莨菪碱
Hyoscyamine
(μg g-1 DW)
T0 1058.072±37.583 a 1083.528±31.056 d 1052.901±35.189 d
T1 1100.436±16.571 ab 915.777±20.900 a 717.403±22.549 a
T2 1108.081±25.848 ab 923.651±17.580 ab 731.083±22.291 ab
T3 1160.567±33.114 bc 961.774±16.661 b 769.983±19.220 b
T4 1194.663±54.229 c 1005.077±26.956 c 830.383±16.479 c
T5 1160.712±22.361 bc 936.849±19.690 ab 764.317±27.984 b
东莨菪碱
Scopolamine
(μg g-1 DW)
T0 705.793±9.434 c 712.340±4.758 c 706.746±6.777 e
T1 608.653±18.251 a 436.066±42.860 a 322.495±13.838 a
T2 639.789±29.007 ab 443.938±22.239 a 342.123±13.148 ab
T3 703.185±23.661 c 491.396±19.175 ab 363.689±24.281 bc
T4 691.481±28.849 c 533.542±44.175 b 425.387±36.488 d
T5 665.574±21.782 bc 482.438±38.057 ab 384.401±18.008 c

图4

UV-B胁迫下不同浓度Ca2+对颠茄根(A)、叶(B)中PMT、H6H、TR I相对基因表达量的影响 标以不同小写字母的柱值在同一个基因不同处理组间差异显著(P < 0.05)。T0组表示未进行任何处理的空白对照, T1组表示UV-B辐射处理12 d, T4组表示UV-B辐射背景下外施6 mmol L-1 Ca2+处理12 d。"

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