Identification of FtsH gene family and functional analysis of CsFtsH31 gene
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摘要:
目的 FtsH基因在植物的抗逆境胁迫中具有重要的作用，揭示茶树FtsH基因在茶树中的功能和表达模式可为茶树耐光氧化和抗高温胁迫改良育种提供理论依据。 方法 通过茶树基因组数据鉴定FtsH基因，对其进行生物信息学分析，筛选出1个对高温敏感的CsFtsH31基因。在烟草中超量表达CsFtsH31基因，通过光氧化和高温处理分析CsFtsH31基因的表达模式。 结果 在茶树基因组中共鉴定到了45个FtsH基因，按照进化关系可将其分成5类，拟南芥和茶树FtsH基因同源性很高。该家族成员分布在茶树不同的染色体上。茶树中 CsFtsH基因在不同组织和不同胁迫条件下存在差异表达，在高温处理后茶树中CsFtsH14、CsFtsH31、CsFtsH34基因随着时间变化呈上升趋势，CsFtsH31对高温尤其敏感；遗传转化CsFtsH31基因获得转CsFtsH31基因烟草，对转CsFtsH31基因烟草进行光氧化和高温胁迫处理，结果发现转基因烟草的叶绿素、可溶性糖含量及超氧化物歧化酶活性均高于野生型，而丙二醛含量低于野生型。 结论 CsFtsH31基因在烟草中表达，提高叶绿素、可溶性糖含量及超氧化物歧化酶活性，降低丙二醛，清除胁迫产生的活性氧，保护膜结构功能，提高烟草的抗光氧化和抗高温胁迫能力。 Abstract:Objective FtsH gene plays an important role in plant stress resistance. Revealing the function and expression pattern of FtsH gene in tea plant can provide a theoretical basis for the improvement of tea plant resistance to photooxidation and high temperature stress. Method The FtsH gene was identified by tea plant genome data, and a CsFtsH31 gene sensitive to high temperature was screened by bioinformatics analysis. The CsFtsH31 gene was overexpressed in tobacco, and the expression patterns of CsFtsH31 gene against photooxidation and high temperature were analyzed. Result A total of 45 FtsH genes were identified in the genome of tea plant, which were divided into five categories according to the evolutionary relationship. The members of this family are distributed on different chromosomes of tea plants. CsFtsH genes in tea plants are differentially expressed in different tissues and stress conditions. After high temperature treatment, CsFtsH14, CsFtsH31 and CsFtsH34 genes in tea plants showed an increasing trend over time, and CsFtsH31 is particularly sensitive to high temperature. The transgenic tobacco with CsFtsH31 gene was obtained by genetic transformation of CsFtsH31 gene. The transgenic tobacco with CsFtsH31 gene was subjected to stress treatment. The results showed that the chlorophyll and soluble sugar content and superoxide dismutase activity of transgenic tobacco were higher than wild type, while malondialdehyde was lower than wild type. Conclusion The CsFtsH31 gene was expressed in tobacco by increasing the content of chlorophyll and soluble sugar, and the activity of superoxide dismutase, reducing the content of malondialdehyde, scavenging the reactive oxygen species produced by stress, protecting the function of membrane structure, and improving the resistance of tobacco to light oxidation and high temperature stress. -
图 1 茶树 FtsH 基因预测保守蛋白基序
Figure 1. Schematic diagram of conserved protein motif predicted by FtsH gene in tea plant
图 2 茶树和拟南芥FtsH基因家族的系统进化树
Figure 2. Phylogenetic tree of FtsH gene family in C. sinensis and A. thaliana
图 3 茶树FtsH基因在不同组织中表达模式
Figure 3. Expression patterns of FtsH gene in different tissues of C. sinensis
图 4 CsFtsH基因在处理条件下表达模式
A：茉莉酸甲酯处理；CK、12 h-MeJA、24 h-MeJA、48 h-MeJA分别表示茉莉酸甲酯处理0、12、24、48 h。B：NaCl处理；CK、12 h-NaCl、24 h-NaCl、48 h-NaCl分别表示盐处理0、12、24、48 h； C：冷处理；CK、CA1、CA3分别表示未适应、完全适应、去适应； D：干旱处理；CK、24 h-PEG、48 h-PEG、72 h-PEG分别表示聚乙二醇处理0、24、48、72h。
Figure 4. Expression pattern of CsFtsH gene under treatment conditions
A: Methyl jasmonate treatment; CK, 12 h-MeJA, 24 h-MeJA, 48 h-MeJA represent methyl jasmonate treatment for 0, 12, 24, 48 h; B: NaCl treatment ; CK, 12 h-NaCl, 24 h-NaCl, 48 h-NaCl represent salt treatment for 0, 12, 24, 48 h ; C:Cold treatment ; CK, CA1, and CA3 represent unadapted, fully adapted, and deadapted, respectively ; D. Drought treatment; CK, 24 h-PEG, 48 h-PEG and 72 h-PEG represent polyethylene glycol treatment for 0, 24, 48, 72 h, respectively.
图 5 高温处理后茶树CsFtsH基因相对表达量
不同小写字母表示同一处理时间不同基因之间差异显著（P＜0.05）。
Figure 5. Relative expression of CsFtsH gene in tea plant after high temperature treatment
Different lowercase letters indicated significant difference at the same treatment time (P＜ 0.05).
图 6 转CsFtsH31基因烟草的遗传转化过程
A：共转化； B：抗性筛选；C：抗性芽生长； D：生根；E：移栽；WT：野生型烟草；1～4：转基因烟草。
Figure 6. Genetic transformation process of CsFtsH31 transgenic tobacco
A: Co-transformation; B: Resistance screening; C: Resistant bud growth; D: Rooting; E: Transplanting; WT: wide type tobacco; 1～4: transgenic tobacco.
图 7 WT与转基因烟草鉴定结果
A：烟草GUS组织化学染色鉴定结果；B：烟草的PCR鉴定结果；M: Marker DL1500；WT：野生型植株; TP1～TP13：转基因烟草植株。
Figure 7. Identification results of WT and transgenic tobacco
A: tobacco GUS histochemical staining results; B: PCR identification results of tobacco; M: Marker DL1500; WT : wild-type plants; TP1–TP13: transgenic tobacco plants.
图 8 转CsFtsH31基因烟草株系基因相对表达量
WT：野生型植株；TP1～TP13：转基因植株。
Figure 8. Expression of CsFtsH31 gene in transgenic tobacco lines
WT: wild-type plants; TP1–TP13: transgenic tobacco plants.
图 9 转基因烟草和野生型烟草抗光氧化表型
Figure 9. Photooxidation resistance phenotype of transgenic tobacco and wild type tobacco
图 10 光氧化处理下CsFtsH31基因相对表达量
Figure 10. The relative expression of CsFtsH31 gene under photooxidation treatment
图 11 光氧化条件下野生型和转基因烟草物质含量
不同小写字母表示同一处理时间不同植株之间差异显著（P＜0.05）。
Figure 11. The substance content of wild-type and transgenic tobacco under photooxidation conditions
Different lowercase letters indicated significant difference among different plants at the same treatment time (P＜0.05).
图 12 高温处理下转基因烟草和野生型烟草表型
Figure 12. Phenotypic characteristicss of transgenic tobacco and wild-type tobacco under high temperature treatment
图 13 高温条件下CsFtsH31基因相对表达量
Figure 13. The relative expression of CsFtsH31 gene under high temperature conditions
图 14 高温处理野生型和转基因烟草物质含量
Figure 14. Substance content of wild-type and transgenic tobacco under high temperature
表 1 荧光定量PCR引物序列
Table 1. Primer sequences for fluorescent quantitative PCR
基因名称
Gene name引物名称
Primer name序列（5′-3′）
Sequence (5′-3′)CsFtsH7 F GATCTGGTGGAAATCCCTTCTT R CCCTTTCGTATGCCTTCTCTAC CsFtsH14 F GAGATCAAGGACGAGAAGAACC R GAGTAAGGACCCAAGCCATAAG CsFtsH31 F CGATGACCGGATAAAGGTGATAG R TCTTCAGTGGGATGTGGAAAC CsFtsH32 F CCTTGGACCCAGCACTTATAC R AATCCTCCAAGTCAACCTCATC CsFtsH33 F TCTGGCGTTCCTTTCCTTTC R TCGACCAATCGCGTCTATTTC CsFtsH34 F CTTTCTGTTGTTGGGCTTCTTC Actin R GAGCATCCAACCCACCTATATC F CAGACCGTATGAGCAAGGAAAT R GTGCTTAGGGATGCAAGGATAG 
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表 2 茶树FtsH基因家族基因的特征
Table 2. Gene characteristics of FtsH gene family in tea
基因
Gene name染色体号
Chr. No.起始位点
Start site/bp终止位点
End site/bp外显子数
No. of exon氨基酸个数
No. of aa相对分子质量
MW/Da等电点
pI亚细胞定位
Subcellular localizationCsFtsH01 Chr7 65213789 65219324 5 923 74458.18 5.95 叶绿体Chloroplast CsFtsH02 Chr11 8563939 8575357 6 1903 73482.17 5.8 叶绿体Chloroplast CsFtsH03 Chr7 65042017 65048440 8 1071 70228.69 6.83 叶绿体Chloroplast CsFtsH04 Contig94 61525 67949 8 1071 70349.93 6.43 叶绿体Chloroplast CsFtsH05 Chr7 64271114 64274398 7 547 69941.07 5.89 叶绿体Chloroplast CsFtsH06 Chr1 165533920 165545204 6 1881 61562.76 8.87 叶绿体Chloroplast CsFtsH07 Chr4 4915196 4918874 5 613 75160.8 6.22 叶绿体Chloroplast CsFtsH08 Chr4 5085616 5089280 5 611 75160.8 6.22 叶绿体Chloroplast CsFtsH09 Contig614 638742 646402 5 1277 76784.53 6.16 叶绿体Chloroplast CsFtsH10 Chr9 158414219 158419184 5 828 76784.53 6.16 叶绿体Chloroplast CsFtsH11 Chr1 165714763 165716089 5 221 33401.76 8.35 叶绿体Chloroplast CsFtsH12 Chr11 65658737 65665118 8 1064 90462.96 6.67 细胞质Cytoplasm；线粒体Mitochondria CsFtsH13 Chr10 124176692 124190387 8 2283 90339.88 8.35 线粒体Mitochondria；细胞核Nucleus CsFtsH14 Contig1031 102459 114466 8 2001 91207.6 6.49 细胞核Nucleus CsFtsH15 Chr4 143895734 143907523 8 1965 91207.6 6.49 细胞核Nucleus CsFtsH16 Chr14 23913349 23923554 7 1701 76689.48 8.19 线粒体Mitochondria CsFtsH17 Chr1 99001591 99024536 17 3824 86023.11 6.11 细胞质Cytoplasm；细胞核Nucleus CsFtsH18 Contig86 451631 471388 17 3293 86050.19 6.17 叶绿体Chloroplast； CsFtsH19 Chr12 118041782 118052606 7 1804 77336.2 7.28 线粒体Mitochondria CsFtsH20 Chr12 117456893 117466022 7 1522 79049.1 6.35 线粒体Mitochondria；细胞核Nucleus CsFtsH21 Chr4 135148277 135159234 13 1826 90008.46 8.84 细胞核Nucleus CsFtsH22 Chr12 157184785 157193617 15 1472 107804 8.55 细胞质Cytoplasm；线粒体Mitochondria CsFtsH23 Chr11 68555164 68565378 17 1702 87978.33 8.97 叶绿体Chloroplast；线粒体Mitochondria CsFtsH24 Chr7 174884414 174903588 17 3196 97408.82 6.46 叶绿体Chloroplast；线粒体Mitochondria CsFtsH25 Chr15 20064510 20075312 11 1800 99166.96 9.36 细胞质Cytoplasm CsFtsH26 Chr1 120253819 120297976 18 7360 96694.01 7.5 线粒体Mitochondria CsFtsH27 Chr5 2362154 2387569 19 4236 117830.5 7.32 细胞质Cytoplasm；线粒体Mitochondria CsFtsH28 Chr15 26955723 26966388 9 1778 47361.01 8.89 细胞质Cytoplasm；细胞核Nucleus CsFtsH29 Contig18 453025 465709 9 2114 47158.76 8.95 细胞质Cytoplasm；细胞核Nucleus CsFtsH30 Chr14 14096902 14101142 9 707 47200.7 8.84 细胞质Cytoplasm；细胞核Nucleus CsFtsH31 Chr15 4795212 4801284 10 1012 47654.97 6.03 细胞质Cytoplasm；细胞核Nucleus CsFtsH32 Chr6 71004311 71010326 10 1003 47429.08 4.94 细胞质Cytoplasm；细胞核Nucleus CsFtsH33 Contig1045 130115 136539 6 1071 47244.84 5.54 细胞质Cytoplasm；细胞核Nucleus CsFtsH34 Chr11 86059724 86070944 6 1870 49573.11 5.91 细胞质Cytoplasm；细胞核Nucleus CsFtsH35 Chr10 109785037 109791165 9 1021 89802.84 5.13 细胞质Cytoplasm；内质网Endoplasmic reticulum CsFtsH36 Chr4 168750629 168756424 9 966 89762.79 5.11 细胞质Cytoplasm；内质网Endoplasmic reticulum CsFtsH37 Contig1101 130919 132805 4 314 54161.11 7.21 细胞核Nucleus CsFtsH38 Chr13 70484157 70491378 8 1204 48009.44 6.7 细胞核Nucleus CsFtsH39 Chr2 115759771 115772281 7 2085 56980.02 6.54 细胞质Cytoplasm CsFtsH40 Chr13 95060923 95073441 22 2086 87951.74 7.86 细胞质Cytoplasm；内质网Endoplasmic reticulum CsFtsH41 Chr7 26348506 26361837 20 2222 81034.76 6.08 内质网Endoplasmic reticulum CsFtsH42 Chr3 76537449 76573876 14 6071 69962.59 6.83 叶绿体Chloroplast；细胞核Nucleus CsFtsH43 Chr7 184741572 184748019 8 1075 70558.86 9.36 叶绿体Chloroplast；细胞质Cytoplasm CsFtsH44 Chr1 218713083 218714729 1 274 57697.36 9.07 叶绿体Chloroplast CsFtsH45 Chr6 80323466 80324930 3 244 53483.53 9.03 叶绿体Chloroplast
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表 3 茶树CsFtsH 基因顺式作用元件
Table 3. Cis-acting elements of CsFtsH gene in tea plant
编号
No.基因名称
Gene name顺式作用元件 Cis-acting elements 光反应元件 环境调控相关元件 基本元件 激素应答相关元件 蛋白代谢调节元件 生长调控元件 L1 L2 E1 E2 E3 A1 H1 H2 H3 H4 P1 G1 G2 CSS0029386.1 CsFtsh01 √ ─ √ √ √ √ √ √ CSS0041542.1 CsFtsh02 √ √ √ √ √ √ √ CSS0004769.1 CsFtsh03 √ √ √ √ √ √ √ CSS0033124.1 CsFtsh04 √ √ √ √ √ √ √ CSS0045434.1 CsFtsh05 √ √ √ √ √ √ √ √ CSS0025086.1 CsFtsh06 √ √ √ CSS0047800.1 CsFtsh07 √ √ √ √ √ √ √ √ √ CSS0022449.1 CsFtsh08 √ √ √ √ √ √ √ √ √ CSS0045969.1 CsFtsh09 √ √ √ √ √ √ √ √ √ √ CSS0013964.1 CsFtsh10 √ √ √ √ √ √ √ √ √ √ CSS0028048.1 CsFtsh11 √ √ √ √ CSS0008560.1 CsFtsh12 √ √ √ √ √ √ CSS0038960.1 CsFtsh13 √ √ √ √ √ √ √ √ √ CSS0020099.1 CsFtsh14 √ √ √ √ √ CSS0033381.1 CsFtsh15 √ √ √ √ √ √ CSS0033670.1 CsFtsh16 √ √ √ √ √ √ CSS0040067.1 CsFtsh17 √ √ √ √ √ √ √ √ √ CSS0014825.1 CsFtsh18 √ √ √ √ √ √ √ √ √ CSS0002059.1 CsFtsh19 √ √ √ √ √ √ CSS0016987.1 CsFtsh20 √ √ √ √ √ √ CSS0008554.1 CsFtsh21 √ √ √ √ √ √ √ CSS0016692.1 CsFtsh22 √ √ √ √ √ √ √ √ √ CSS0046009.1 CsFtsh23 √ √ √ √ √ √ √ CSS0034081.1 CsFtsh24 √ √ √ √ √ CSS0004299.1 CsFtsh25 √ √ √ √ √ √ √ √ √ √ CSS0046503.1 CsFtsh26 √ √ √ √ √ √ CSS0042976.1 CsFtsh27 √ √ √ √ √ CSS0028230.1 CsFtsh28 √ √ √ √ CSS0027170.1 CsFtsh29 √ √ √ CSS0029520.1 CsFtsh30 √ √ √ CSS0048047.1 CsFtsh31 √ √ √ √ √ √ CSS0018374.1 CsFtsh32 √ √ √ √ √ √ CSS0043566.1 CsFtsh33 √ √ √ √ √ √ √ √ CSS0047454.1 CsFtsh34 √ √ √ √ √ CSS0008341.1 CsFtsh35 √ √ √ √ √ CSS0041444.1 CsFtsh36 √ √ √ √ √ √ √ √ CSS0000928.1 CsFtsh37 √ √ √ √ √ √ CSS0009578.1 CsFtsh38 √ √ √ √ √ √ CSS0004467.1 CsFtsh39 √ √ √ √ √ √ √ √ CSS0001611.1 CsFtsh40 √ √ √ √ √ √ √ CSS0027753.2 CsFtsh41 √ √ √ √ √ CSS0022555.1 CsFtsh42 √ √ √ √ √ CSS0028493.1 CsFtsh43 √ √ √ √ √ √ √ CSS0014817.1 CsFtsh44 √ √ √ √ √ √ CSS0033189.1 CsFtsH45 √ √ √ √ "L1、L2表示光反应元件Light、P-box；E1~E3表示环境调控元件TGA-element、MBS、LTR；A1表示基本元件CAAT-box；H1~H4表示激素应答相关元件ABRE、TGA-element、CGTCA-motif、TCA-element；P1表示蛋白代谢调节元件O2-site；G1、G2表示生长调控元件Circadian、CAT-box。
L1, L2 represent the light reaction element Light and P-box; E1– E3 represent the environmental regulatory element TGA-element, MBS and LTR; A1 represents the basic element CAAT-box; H1–H4 represent ABRE, TGA-element, CGTCA-motif and TCA-element; P1 represents the protein metabolic regulatory element O2-site; G1 and G2 represent the growth regulatory elements Circadian and CAT-box."
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