蒙古冰草肉桂醇脱氢酶基因序列鉴定及功能分析

doi:10.11983/CBB23109

植物学报 ›› 2024, Vol. 59 ›› Issue (2): 204-216.DOI: 10.11983/CBB23109

蒙古冰草肉桂醇脱氢酶基因序列鉴定及功能分析

王贺萍¹^,², 孙震², 刘雨辰², 苏彦龙², 杜锦瑜²^,⁵, 赵彦⁵, 赵竑博¹, 王召明⁶, 苑峰⁶, 刘亚玲⁶, 吴振映²^,³^,⁴, 何峰²^,³^,⁴^,^*(), 付春祥²^,³^,⁴^,^*()

¹华南农业大学园艺学院, 广州 510642
²中国科学院青岛生物能源与过程研究所, 青岛 266101
³山东能源研究院, 青岛 266101
⁴青岛新能源山东省实验室, 青岛 266101
⁵内蒙古农业大学草原与资源环境学院/农业农村部饲草栽培、加工与高效利用重点实验室/草地资源教育部重点实验室,呼和浩特 010018
⁶内蒙古自治区牧草育种与良种繁育重点实验室, 呼和浩特 010018

收稿日期:2023-08-09 接受日期:2023-12-19 出版日期:2024-03-01 发布日期:2024-01-24
通讯作者: * E-mail: hefeng@qibebt.ac.cn; fucx@qibebt.ac.cn
基金资助:
内蒙古自治区种业科技创新重大示范工程“揭榜挂帅”项目(2022JBGS0014);中央引导地方科技发展资金(2022ZY006);呼和浩特市科技创新领域人才项目(“政产学研推用银”创新联合体项目)(2022RC-联合体-2);山东能源研究院科研创新基金(SEII202142)

Sequence Identification and Functional Analysis of Cinnamyl Alcohol Dehydrogenase Gene from Agropyron mongolicum

Heping Wang¹^,², Zhen Sun², Yuchen Liu², Yanlong Su², Jinyu Du²^,⁵, Yan Zhao⁵, Hongbo Zhao¹, Zhaoming Wang⁶, Feng Yuan⁶, Yaling Liu⁶, Zhenying Wu²^,³^,⁴, Feng He²^,³^,⁴^,^*(), Chunxiang Fu²^,³^,⁴^,^*()

¹College of Horticulture, South China Agricultural University, Guangzhou 510642, China
²Qingdao Institute of Bioenergy and Bioprocess Technology, Chinese Academy of Sciences, Qingdao 266101, China
³Shandong Energy Institute, Qingdao 266101, China
⁴Qingdao New Energy Shandong Laboratory, Qingdao 266101, China
⁵Key Laboratory of Grassland Resources, Ministry of Education/Key Laboratory of Forage Cultivation, Processing and Efficient Utilization of Ministry of Agriculture and Rural Affairs/College of Grassland, Resource and Environmental Science, Inner Mongolia Agricultural University, Hohhot 010018, China
⁶Key Laboratory of Forage Breeding and Seed Production in Inner Mongolia Autonomous Region, Hohhot 010018, China

Received:2023-08-09 Accepted:2023-12-19 Online:2024-03-01 Published:2024-01-24
Contact: * E-mail: hefeng@qibebt.ac.cn; fucx@qibebt.ac.cn

摘要/Abstract

摘要： 肉桂醇脱氢酶(CAD)作为植物次生代谢尤其是木质素生物合成过程的关键酶, 在调控植物生长发育和抵御生物/非生物胁迫等过程中发挥关键作用。蒙古冰草(即沙芦草(Agropyron mongolicum))耐旱耐寒, 在我国北方荒漠草原区域广泛分布。为探讨CAD基因在蒙古冰草木质素合成和非生物胁迫抗性中的作用, 从蒙古冰草全长转录组数据中筛选并克隆到1个CAD基因, 序列长度1 083 bp, 命名为AmCAD。该基因编码361个氨基酸残基, 同源序列比对发现蛋白质序列保守区域含有2个Zn²⁺结合基序和NADP(H)辅因子结合基序, 属于典型的CAD蛋白, 且三维结构与AtCAD5相似。AmCAD在茎秆中高表达, 对AmCAD重组蛋白的酶学性质分析表明, 该蛋白对不同肉桂醛类底物均具有很强的催化能力, 其中对松柏醛和芥子醛的底物亲和力更强。用不同浓度甘露醇模拟干旱胁迫, 蒙古冰草AmCAD基因表达受到显著诱导。研究结果表明, AmCAD在蒙古冰草木质素合成和干旱胁迫抗性中发挥重要作用, 可为提高蒙古冰草品质和抗逆性分子育种提供有价值的基因资源。

关键词: 蒙古冰草, 肉桂醇脱氢酶, 木质素, 抗旱性

Abstract: As an essential enzyme in plant secondary metabolism, cinnamyl alcohol dehydrogenase (CAD) plays a key role in regulating plant growth and development, as well as biological/abiotic stress resistance. Agropyron mongolicum is a traditional forage grass widely distributed in the desert grassland areas of northern China, which exhibited high tolerance to drought and cold stresses. To explore the role of cinnamyl alcohol dehydrogenase in A. mongolicum, in this study, a CAD gene was identified from the full-length transcriptome data of A. mongolicum and subsequentially analyzed in vitro. The 1 083 bp coding sequence of AmCAD encodes 361 amino acids, which has typical conserved CAD region containing two Zn²⁺binding motifs and NADP(H) cofactor binding motifs, belongs to the typical CAD protein, and its three-dimensional structure is similar to AtCAD5. AmCAD is highly expressed in the stem. The AmCAD recombinant protein showed a robust catalytic ability to different cinnamaldehyde substrates, with the highest substrate affinity of coniferyl aldehyde and sinapaldehyde. Under drought stress condition, the expression level of AmCAD was significantly induced, indicating a potential function of this gene in stress tolerance. The experimental results indicate that AmCAD may play an important role in lignin biosynthesis and drought stress tolerance in A. mongolicum. Our research provided potentially valuable genetic resources for molecular breeding of A. mongolicum to improve biomass quality and stress resistance.

Key words: Agropyron mongolicum, cinnamyl alcohol dehydrogenase, lignin, drought resistance

王贺萍, 孙震, 刘雨辰, 苏彦龙, 杜锦瑜, 赵彦, 赵竑博, 王召明, 苑峰, 刘亚玲, 吴振映, 何峰, 付春祥. 蒙古冰草肉桂醇脱氢酶基因序列鉴定及功能分析. 植物学报, 2024, 59(2): 204-216.

Heping Wang, Zhen Sun, Yuchen Liu, Yanlong Su, Jinyu Du, Yan Zhao, Hongbo Zhao, Zhaoming Wang, Feng Yuan, Yaling Liu, Zhenying Wu, Feng He, Chunxiang Fu. Sequence Identification and Functional Analysis of Cinnamyl Alcohol Dehydrogenase Gene from Agropyron mongolicum. Chinese Bulletin of Botany, 2024, 59(2): 204-216.

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https://www.chinbullbotany.com/CN/Y2024/V59/I2/204

图/表 8

表1 蒙古冰草AmCAD蛋白理化性质分析

Table 1 Analysis of physicochemical properties of AmCAD proteins from Agropyron mongolicum

Sequence ID	Number of amino acid	Molecular weight (Da)	Theoretical pI	Instability index	Aliphatic index	Grand average of hydropathicity
PB.70108.1	360	38591.48	5.87	23.05	90.33	0.025
PB.82665.1	354	38437.86	6.66	24.94	84.75	-0.098
PB.80567.1	358	38685.52	6.60	30.79	89.27	0.012
PB.80906.1	355	38256.89	5.68	31.84	88.42	0.037
PB.62308.1	269	28625.35	6.31	31.41	92.45	0.166
PB.17482.1	344	36802.76	5.86	31.22	86.13	0.103
PB.52032.1	373	38747.35	6.53	24.05	91.58	0.136
PB.47633.2	421	44353.00	6.42	31.04	84.28	0.092
PB.55683.1	421	44338.97	6.42	29.60	83.80	0.088
PB.83678.1	356	37359.82	5.40	25.49	92.25	0.146
PB.84043.1	356	37036.55	5.84	26.83	93.88	0.211
PB.84184.1	356	37019.73	6.08	23.61	92.25	0.195

图1 蒙古冰草AmCAD蛋白的基序分析

Figure 1 Motif analysis of AmCAD protein in Agropyron mongolicum

图2 蒙古冰草与拟南芥、水稻和二穗短柄草CAD的系统进化树红色五角星代表可能参与木质素合成的蒙古冰草CAD蛋白; 粉色点代表二穗短柄草CAD蛋白; 绿色点代表拟南芥CAD蛋白; 蓝色点代表水稻CAD蛋白。

Figure 2 Phylogenetic tree of CADs in Agropyron mongolicum, Arabidopsis thaliana, Oryza sativa, and Brachypodium distachyon The red pentagram represents the functional CAD protein of A. mongolicum that may be involved in lignin synthesis; the pink dots represent the CAD protein of B. distachyon; the green dots represent the CAD protein of A. thaliana; and the blue dots represent the CAD protein of O. sativa.

图3 蒙古冰草AmCAD基因扩增和蛋白结构分析 (A) AmCAD电泳图(M: DNA ladder; 1: 扩增序列); (B) 蒙古冰草与其它物种CAD氨基酸序列比对(构成底物结合口袋的残基用三角形(▲)标记); (C) AmCAD蛋白质三维结构模型和底物结合口袋(底部亚基的核苷酸结合和催化结构域分别呈黄色和蓝色, 在上部亚基中的对应结构分别以绿色和深蓝色显示, 锌离子用灰色球体表示)

Figure 3 Amplification of Agropyron mongolicum AmCAD and protein structure analysis (A) AmCAD gel map (M: DNA ladder; 1: Amplification sequence); (B) Comparison of CAD amino acid sequences among A. mongolicum and other species (the residue forming the substrate binding pocket is marked with a triangle (▲)); (C) Three-dimensional protein structure model and substrate binding pocket of AmCAD (the nucleotide binding and catalytic domains of the bottom subunit are colored yellow and blue, respectively, the corresponding compounds in the upper subunit are colored green and dark blue, while the zinc ions are indicated by gray spheres)

图4 AmCAD重组蛋白在大肠杆菌中的异源表达(A)及不同温度(B)和pH (C)下的酶学性质 1: 蛋白Marker; 2: 上清液; 3: 沉淀; 4: 流出液; 5: 纯化蛋白

Figure 4 Heterologous expression (A) and enzymatic properties of AmCAD recombinant protein in Escherichia coli at different temperatures (B) and pH (C) 1: Marker; 2: Supernatant; 3: Precipitant; 4: Effluent; 5: Purified protein

表2 AmCAD重组蛋白催化不同底物的酶动力学特性

Table 2 Enzyme kinetic characteristics of different substrates catalyzed by recombinant AmCAD

Substract	K_m (μmol·L^-1)	V_max(μmol·L^-1·s^-1)	K_cat (·s⁻¹)	K_cat/K_m(μmol·L^-1·s^-1)
Coniferylaldehyde	8.94±0.15	6.87±0.008	441.37±0.53	49.35±0.90
Sinapaldehyde	10.35±2.14	7.18±0.43	461.51±27.94	45.26±6.69
p-coumaraldehyde	35.35±3.74	15.21±1.60	977.60±102.97	27.65±0.01

图5 AmCAD在蒙古冰草不同组织(A)和茎节部位(B)的表达量

Figure 5 Expression levels of AmCAD in different tissues (A) and internodes (B) of Agropyron mongolicum

图6 AmCAD基因在不同处理时间及不同浓度甘露醇模拟干旱胁迫下的表达分析 (A) 75 mmol∙L-1甘露醇处理; (B) 150 mmol∙L-1甘露醇处理; (C) 250 mmol∙L-1甘露醇处理。* P<0.05; ** P<0.01; *** P<0.001

Figure 6 Expression analysis of AmCAD gene under drought stress at different time points and mannitol concentrations (A) 75 mmol∙L-1 mannitol treatment; (B) 150 mmol∙L-1 mannitol treatment; (C) 250 mmol∙L-1 mannitol treatment. * P<0.05; ** P<0.01; *** P<0.001

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蒙古冰草肉桂醇脱氢酶基因序列鉴定及功能分析

Sequence Identification and Functional Analysis of Cinnamyl Alcohol Dehydrogenase Gene from Agropyron mongolicum

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