江西铅山红芽芋超低温疗法脱毒苗的转录组分析

尹明华; 张艺欣; 邓雨晴; 吴洪婷; 陈婷; 郭淑贞; 杨于萱

doi:10.19303/j.issn.1008-0384.2020.10.002

江西铅山红芽芋超低温疗法脱毒苗的转录组分析

doi: 10.19303/j.issn.1008-0384.2020.10.002

尹明华^{1, 2, 3,},
张艺欣¹,
邓雨晴¹,
吴洪婷¹,
陈婷¹,
郭淑贞¹,
杨于萱¹

1.
上饶师范学院生命科学学院，江西　上饶　334001
2.
上饶市药食同源植物资源保护与利用重点实验室，江西　上饶　334001
3.
上饶农业技术创新研究院，江西　上饶　334001

基金项目: 国家自然科学基金项目（31860084、32060092）；上饶师范学院校级自选课题（202031）；江西省大学生创新创业训练计划项目（S202010416014）

详细信息

作者简介:
尹明华（1973−），女，硕士，副教授，主要从事植物生物技术方面研究（E-mail：yinminghua04@163.com）

中图分类号: S 532
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出版历程
- 收稿日期: 2020-03-10
- 修回日期: 2020-07-15
- 刊出日期: 2020-10-28

Transcriptome Analysis on Cryotherapy-treated Virus-free Plantlets of Red Bud Taro at Yanshan, Jiangxi Province

1.
College of Life Sciences, Shangrao Normal University, Shangrao, Jiangxi　334001, China
2.
Key Laboratory of Protection and Utilization of Medicinal and Edible Plant Resources in Shangrao City, Shangrao, Jiangxi　334001, China
3.
Shangrao Agricultural Technology Innovation Research Institute, Shangrao, Jiangxi　334001, China

摘要

摘要: 目的探究江西铅山红芽芋对超低温疗法脱毒的转录组响应机制，为江西铅山红芽芋超低温疗法脱毒苗的规模化应用奠定理论依据。方法以江西铅山红芽芋超低温疗法脱毒苗（VF组）和江西铅山红芽芋超低温疗法带毒苗（V组）试验材料进行转录组比较分析。结果 VF组Clean reads为42 406 188，GC含量为54.09%；V组Clean reads为46 818 060，GC含量为50.36%。VF组和V组表达量FPKM的对数值在0～2，表达量密度在0～0.8。VF组和V组表达的共有基因数为23 820，V组单独表达的基因数为10 298，VF组单独表达的基因数为4 477。VF组和V组表达量的相关系数为0.287，样本间相关性较差。VF组和V组共产生差异表达基因5 282 个，与V组比较，VF组上调基因3 011，下调基因2 271。GO 富集分析显示，差异基因主要注释到过氧化氢分解过程、过氧化氢代谢过程、一元羧酸生物合成过程、多糖分解代谢过程、金属离子输运、细胞外区、细胞壁、外部封装结构、膜的固有成分、质膜固有成分、核酸结合转录因子活性、序列特异性DNA结合转录因子活性、单加氧酶活性、铁离子结合、血红素结合等功能。KEGG 富集分析显示，差异基因主要注释到苯丙酸生物合成、类黄酮生物合成、二苯乙烯类和二芳基庚烷类以及姜辣素的生物合成、MAPK信号通路-植物、淀粉和蔗糖代谢、酪氨酸代谢、植物激素信号转导、戊糖和葡萄糖醛酸的相互转化、过氧化物酶体、α-亚麻酸代谢、苯丙氨酸代谢、氰胺酸代谢、类胡萝卜素生物合成、异喹啉生物碱的生物合成、泛醌和其他萜类醌的生物合成、甘氨酸、丝氨酸和苏氨酸代谢、植物昼夜节律、果糖和甘露糖代谢、半乳糖代谢、氨基糖和核苷酸糖代谢等途径。结论 F-box家族蛋白、脱落酸受体PYR1、乙烯受体2、生长素响应因子1、BZIP转录因子家族蛋白、过氧化物酶、过氧化氢酶1、超氧化物歧化酶[Cu-Zn]、植物抗病反应蛋白、抗病蛋白、抗病蛋白RPS2、抗病蛋白RPS5、抗病蛋白RGA2、抗病反应蛋白、转座子TNT的逆转录病毒相关Pol多蛋白、逆转录病毒相关Pol多聚蛋白系1、类烟草病毒增殖蛋白3、烟草病毒增殖蛋白1、转座子RE1的逆转录病毒相关Pol多蛋白、蔗糖合成酶等基因是江西铅山红芽芋的超低温疗法脱毒的主要响应基因。
- 江西铅山红芽芋 /
- 超低温疗法脱毒苗 /
- 超低温疗法带毒苗 /
- 转录组分析
Abstract: Objective Mechanism of red bud taro in response to cryotherapy was studied by transcriptome analysis for extended application of the virus-free seedlings at Yanshan, Jiangxi. Method Transcriptomes of virus-free (VF) and virus-carrying (V) red bud taro plantlets that had been cryo-therapeutically treated were analyzed and compared. Result In the VF plantlets, the clean reads were 42 406 188 with a GC content of 54.09%; while, in the V plantlets, the clean reads were 46 818 060 with a GC content of 50.36%. For either group, the FPKM was between 0-2, and the expression density between 0-0.8. There were 23 820 genes commonly expressed in both VF and V, while 4 477 independently expressed in VF, and 10 298 in V. There was a low correlation between VF and V with a coefficient of 0.287, and were 5 282 differentially expressed genes (DEG) in them. Compared to V, VF had 3 011 genes were up-regulated and 2 271 down-regulated. The Go enrichment analysis showed that the differential genes were annotated mainly in the processes of hydrogen peroxide catabolism, hydrogen peroxide metabolism, monocarboxylic acid biosynthesis, and polysaccharide catabolism, metal ion transport, extracellular region, cell wall, external encapsulating structure, intrinsic components of membrane and plasma membrane as well as the activities of nucleic acid binding transcription factor, transcription factor, sequence-specific DNA binding, monooxygenase, iron ion binding, heme binding, etc. The KEGG enrichment analysis indicated that the differential genes were annotated largely to the biosynthesis of phenylpropanoid, flavonoid, stilbenoid, diarylheptanoid, gingerol, carotenoid, isoquinoline alkaloid, ubiquinone, and other terpenoid-quinones, the metabolisms of starch, sucrose, tyrosine, alpha-Linolenic acid, phenylalanine, cyanoamino acid, glycine, serine, threonine, fructose, mannose, galactose, amino sugar, and nucleotide sugar, and the pathways of MAPK signaling-plant, plant hormone signal transduction, pentose and glucuronate interconversions, peroxisome, circadian rhythm-plant, and others, Conclusion The genes of F-box family protein, abscisic acid receptor PYR1, ethylene receptor 2, auxin response factor 1, BZIP transcription factor family protein, peroxidase, catalase 1, superoxide dismutase [Cu-Zn], plant disease resistance response protein, disease resistance protein, disease resistance protein RPS2, disease resistance protein RPS5, disease resistance protein RGA2, disease resistance response protein, retrovirus-related Pol polyprotein from transposon TNT, retrovirus-related Pol polyprotein LINE-1, tobamovirus multiplication protein 3-like, tobamovirus multiplication protein 1, retrovirus-related Pol polyprotein from transposon RE1, sucrose synthase, etc. were the predominant genes in the virus-free red bud taro plantlets that responded to the cryotherapy.
- Red bud taro in Jiangxi Province /
- cryotherapy-treated virus-free plantlet /
- cryotherapy-treated virus-carrying plantlet /
- transcriptome analysis

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图 1 VF组和V组表达量分布盒形图（A）和密度图（B）

Figure 1. Box diagram (A) and density diagram (B) on gene expression distributions of VF and V plantlets

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图 2 VF组和V组表达量样本间Venn图

Figure 2. Venn diagram between VF and V plantlets

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图 3 VF组和V组表达量样本间相关性热图

Figure 3. Thermogram on correlation between VF and V plantlets

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图 4 VF组和V组表达量差异火山图（A）和表达量差异散点图（B）

Figure 4. Volcano map (left) and scatter map (right) on gene expression difference between VF and V plantlets

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图 5 VF组和V组差异表达基因GO富集分析柱形图（A）和气泡图（B）

Figure 5. Column map (A) and bubble map (B) of Go enrichment analysis on differentially expressed genes of VF and V plantlets

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图 6 差异表达基因KEGG富集分析柱形图（A）和气泡图（B）

Figure 6. Column chart (A) and bubble chart (B) of KEGG enrichment analysis on differentially expressed genes

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表 1 qRT-PCR引物序列

Table 1. Primers designed for qRT-PCR

基因 Gene	方向 Direction	引物序列 Primer sequence（5′-3′）	大小 Size/bp	TM值 TM value
抗病反应蛋白 Disease resistance response protein	F	TTCGCGACGAAAGGTAAAACGA	105	55
抗病反应蛋白 Disease resistance response protein	R	CACCCACAGGATGGACCAGAGG	105	55
抗病蛋白RGA2 Disease resistance protein RGA2	F	CGGTTCAGAAGGGAGCCTATTGT	199	53.3
抗病蛋白RGA2 Disease resistance protein RGA2	R	AAACTCTTATACCTTTGGGGGGC	199	53.3
乙烯受体2 Ethylene receptor 2	F	TCCCTTTTGACCTTGTTCTTTTG	137	52
乙烯受体2 Ethylene receptor 2	R	TACATTCTCCCGTGTACTTGCAG	137	52
过氧化氢酶1 Catalase 1	F	CAGTGTCAAGACACGGTCGCATG	198	53.8
过氧化氢酶1 Catalase 1	R	CTAGTAACCGCTAGGGCCGTTGG	198	53.8
蔗糖合成酶 Sucrose synthase	F	GCCCCGTCTTATTCATACCCTCG	173	54.6
蔗糖合成酶 Sucrose synthase	R	CATGCTTTTCCCATTTTGTCCTCA	173	54.6
抗病蛋白RPS5 Disease resistance protein RPS5	F	GTCCCAACTTATCCGTCATTTCT	168	51.4
抗病蛋白RPS5 Disease resistance protein RPS5	R	TAAAGGAGCTGCCACTCACAGTC	168	51.4
抗病蛋白 Disease resistance protein	F	TGTGTCTCCCTCCAGTGCTCA	145	53.6
抗病蛋白 Disease resistance protein	R	TTTCTCGCCCCCCCTGTTCA	145	53.6
类烟草病毒增殖蛋白3 Tobamovirus multiplication protein 3-like	F	GGTGTCTCGTTGTTTGCTGCTC	103	52.1
类烟草病毒增殖蛋白3 Tobamovirus multiplication protein 3-like	R	GTTTCTTGCGTCGCCCTTTC	103	52.1
植物抗病反应蛋白 Plant disease resistance response protein	F	TGCCCTGAGCGTTCCCTAC	178	58.3
植物抗病反应蛋白 Plant disease resistance response protein	R	ACCCGCCTCCACTTCTTCC	178	58.3
F-box家族蛋白 F-box family protein	F	CGCCAAGAGGGAAATGAAACC	195	58.5
F-box家族蛋白 F-box family protein	R	CGGGAACATGGAGCAGTGGTA	195	58.5
超氧化物歧化酶[Cu-Zn] Superoxide dismutase [Cu-Zn]	F	CTGAGGCAACGATTGTGGATA	130	53.2
超氧化物歧化酶[Cu-Zn] Superoxide dismutase [Cu-Zn]	R	CCCGTGCTAAGGCTAAGTTCAT	130	53.2
逆转录病毒相关Pol多聚蛋白系-1 Retrovirus-related Pol polyprotein line-1	F	CCCTTCTTTGTTAATGCCCACCA	156	49.9
逆转录病毒相关Pol多聚蛋白系-1 Retrovirus-related Pol polyprotein line-1	R	CGTCCTTAATGCACGGAAGCA	156	49.9
转座子RE1的逆转录病毒相关Pol多蛋白 Retrovirus-related Pol polyprotein from transposon RE1	F	GTCCAGATTGAAGTCGCTCCC	117	53.5
	R	CCTTGTACTGTGACAACAATGCC	117	53.5
转座子TNT逆转录病毒相关Pol多蛋白 Retrovirus-related Pol polyprotein from transposon TNT	F	GATAGACCATCAGGCACTTTAGG	151	48.5
	R	CTAAGTCATTCTTGGGACACTGTAA	151	48.5
BZIP转录因子家族蛋白 BZIP transcription factor family protein	F	CATCGCCGAGCACTTGCACCGTCTC	95	55.6
BZIP转录因子家族蛋白 BZIP transcription factor family protein	R	GCTTCTGGCATCTTTACCACTTTC	95	55.6
过氧化物酶 Peroxidase	F	CCAACTCCCAGGACTTCTTCTTCC	136	57.9
过氧化物酶 Peroxidase	R	CTGGCTGGCTAGTGGTGCTTGTT	136	57.9
脱落酸受体PYR1 Abscisic acid receptor PYR1	F	CTCCATTTCGCCCTGATTCCATT	139	54.6
脱落酸受体PYR1 Abscisic acid receptor PYR1	R	TCAAACATCCCTTTCACCATTCCTAT	139	54.6
抗病蛋白RPS2 Disease resistance protein RPS2	F	ATCCATTCCTTGCCAGATGAC	115	53.4
抗病蛋白RPS2 Disease resistance protein RPS2	R	ACCTTATTAACCGCAGCACCA	115	53.4
生长素反应因子1 Auxin response factor 1	F	CTTTCACGCCCAATCGACCAC	169	57.6
生长素反应因子1 Auxin response factor 1	R	TCGGCTTCTTGCTTTCTGCTGTC	169	57.6
烟草病毒增殖蛋白1 Tobamovirus multiplication protein 1	F	CTGTGGGAAGACTTCTTGCCTGAT	108	50.9
烟草病毒增殖蛋白1 Tobamovirus multiplication protein 1	R	TCGCTTTGGTATTATTGGACCTG	108	50.9
甘油醛-3-磷酸脱氢酶 GAPDH （内参 Reference gene）	F	ATCAAGCCCTCAACAATGCCAAA	179	54.8
甘油醛-3-磷酸脱氢酶 GAPDH （内参 Reference gene）	R	GCCAAGAAGGTCGTCATCTCAGC	179	54.8

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表 2 V组和VF组测序数据统计

Table 2. Statistical sequencing data on V and VF plantlets

样本 Sample	带毒苗 V	脱毒苗 VF
原始序列 Raw reads	47 418 842	43 025 444
原始总碱基数 Raw bases	7 160 245 142	6 496 842 044
过滤后序列 Clean reads	46 818 060	42 406 188
质控后总碱基数 Clean bases	6 964 653 736	6 305 931 529
测序错误率 Error rate/%	0.023 1	0.024 6
大于20的碱基数占总碱基的百分比 Q20/%	98.83	98.26
大于30的碱基数占总碱基的百分比 Q30/%	96.08	94.50
G、C碱基的总数量占总碱基数量的百分比 GC content/%	50.36	54.09

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表 3 测序数据与组装结果比对统计

Table 3. Comparison on sequencing data and assembly results

样本 Sample	过滤后测序数据的条数 Clean reads	能比对到组装转录本上的 Clean reads数 Mapped reads	能定位到组装转录本上的Clean reads所占百分比 Mapped ratio/%
脱毒苗VF	42 406 188	34 683 956	81.79
带毒苗V	46 818 060	38 036 872	81.24

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表 4 VF组和V组表达量差异统计（前10）

Table 4. Top 10 statistical expression differences between VF and V plantlets

基因编号 Gene_ID	差异表达倍数 FC（VF/V）	差异表达倍数以2为底的对数值 log2FC （VF/V）	P值 P value	矫正后 P值 P adjust	显著性 Significance	调节 Regulate	带毒苗表达量 Expression in V	脱毒苗表达量 Expression in VF	NR描述 NR description
TRINITY_DN56_c0_g1	5.15×10⁻⁶	−17.57	5.30×10⁻⁴¹	1.65×10⁻³⁶	是 Yes	下调 Down	33445.52	0.12	假设蛋白质 Hypothetical protein
TRINITY_DN56_c1_g1	5.39×10⁻⁶	−17.50	8.29×10⁻⁴¹	1.65×10⁻³⁶	是 Yes	下调 Down	2710.44	0.01	假设蛋白质 MIMGU_mgv1a026582mg, hypothetical protein MIMGU_mgv1a026582mg
TRINITY_DN388_c0_g1	1.36×10⁻⁵	−16.16	4.43×10⁻⁴⁰	5.88×10⁻³⁶	是 Yes	下调 Down	37524.60	0.40	假设蛋白质 Hypothetical protein
TRINITY_DN56_c0_g2	2.62×10⁻⁵	−15.22	7.54×10⁻⁴⁰	7.52×10⁻³⁶	是Yes	下调 Down	24571.29	0.53	假设蛋白质 Hypothetical protein
TRINITY_DN1368_c0_g1	2.82×10⁻⁵	−15.11	8.27×10⁻³⁹	6.59×10⁻³⁵	是 Yes	下调 Down	16978.15	0.39	未注释 No
TRINITY_DN863_c0_g2	2.14×10⁻⁵	−15.51	4.02×10⁻³⁸	2.67×10⁻³⁴	是 Yes	下调 Down	2861.27	0.05	假设蛋白质 MIMGU_mgv1a026582mg hypothetical protein MIMGU_mgv1a026582mg
TRINITY_DN1333_c0_g1	2.42×10⁻⁵	−15.33	1.41×10⁻³⁷	8.01×10⁻³⁴	是 Yes	下调 Down	8296.82	0.16	未注释 No
TRINITY_DN893_c0_g2	2.93×10⁻⁶	−18.38	3.72×10⁻³⁵	1.86×10⁻³¹	是 Yes	下调 Down	11335.30	0.00	假设蛋白质 Hypothetical protein
TRINITY_DN4426_c0_g1	8.51×10⁻⁵	−13.52	7.01×10⁻³⁵	3.11×10⁻³¹	是 Yes	下调 Down	31665.60	4.91	假设蛋白质 Hypothetical protein
TRINITY_DN6139_c0_g1	6.48×10⁻⁵	−13.91	4.83×10⁻³⁴	1.93×10⁻³⁰	是 Yes	下调 Down	17471.00	0.87	外壳蛋白 Coat protein

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表 5 部分差异表达基因GO富集结果

Table 5. GO enrichment results of some differentially expressed genes

富集到该GO term的基因转录本数目 Number of genes enriched to the GO term	GO Term对应的编号 ID Corresponding to Go term	GO三大分类 Three categories of GO	GO功能描述 Go function description	该GO在目标基因集中占有的比例 The proportion of the GO in the target gene set/%	该GO在背景基因转录本中占有的比例 The proportion of the GO in the background gene/%	未经校正的P值 P value uncorrected	校正后的P值 P value corrected
37	GO:0042744	生物过程 BP	过氧化氢分解过程 Hydrogen peroxide catabolic process	0.62	0.35	0.00	0.00
37	GO:0042743	生物过程 BP	过氧化氢代谢过程 Hydrogen peroxide metabolic process	0.62	0.35	0.00	0.00
65	GO:0072330	生物过程 BP	一元羧酸生物合成工艺 Monocarboxylic acid biosynthetic process	1.09	0.75	0.00	0.02
48	GO:0000272	生物过程 BP	多糖分解代谢过程 Polysaccharide catabolic process	0.80	0.54	0.00	0.04
54	GO:0030001	生物过程 BP	金属离子输运 Metal ion transport	0.91	0.62	0.00	0.04
128	GO:0005576	细胞组分 CC	细胞外区 Extracellular region	2.15	1.45	0.00	0.00
73	GO:0005618	细胞组分 CC	细胞壁 Cell wall	1.23	0.82	0.00	0.00
73	GO:0030312	细胞组分 CC	外部封装结构 External encapsulating structure	1.23	0.82	0.00	0.00
2136	GO:0031224	细胞组分 CC	膜的固有成分 Intrinsic component of membrane	35.88	34.47	0.00	0.15
37	GO:0031226	细胞组分 CC	质膜固有成分 Intrinsic component of plasma membrane	0.62	0.43	0.00	0.18
209	GO:0001071	分子功能 MF	核酸结合转录因子活性 Nucleic acid binding transcription factor activity	3.51	2.48	0.00	0.00
209	GO:0003700	分子功能 MF	序列特异性DNA结合转录因子活性 Transcription factor activity, sequence-specific DNA binding	3.51	0.48	0.00	0.00
118	GO:0004497	分子功能 MF	单加氧酶活性 Monooxygenase activity	1.98	1.27	0.00	0.00
137	GO:0005506	分子功能 MF	铁离子结合 Iron ion binding	2.30	1.54	0.00	0.00
147	GO:0020037	分子功能 MF	血红素结合 Heme binding	2.47	1.69	0.00	0.00

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表 6 差异表达基因KEGG富集部分分析

Table 6. Partial results of analysis on KEGG enrichment of differentially expressed genes

基因数目 Gene number	通路编号 Pathway ID	描述 Description	该KEGG在目标基因集中占有的比例 The proportionof KEGG in target gene concentration/%	该KEGG在背景中占有的比例 The proportion of the KEGG in the background/%	未经校正的P值 P value uncorrected	校正后的P值 P value corrected
78	map00940	苯丙酸生物合成 Phenylpropanoid biosynthesis	2.71	1.36	0.00	0.00
26	map00941	类黄酮生物合成 Flavonoid biosynthesis	0.90	0.38	0.00	0.00
17	map00945	二苯乙烯类、二芳基庚烷类和姜辣素的生物合成 Stilbenoid, diarylheptanoid and gingerol biosynthesis	0.59	0.26	0.00	0.00
71	map04016	MAPK信号通路-植物 MAPK signaling pathway-plant	2.47	1.67	0.00	0.00
63	map00500	淀粉和蔗糖代谢 Starch and sucrose metabolism	2.19	1.52	0.00	0.01
26	map00350	酪氨酸代谢 Tyrosine metabolism	0.90	0.52	0.00	0.01
92	map04075	植物激素信号转导 Plant hormone signal transduction	3.20	2.42	0.00	0.01
26	map00040	戊糖和葡萄糖醛酸的相互转化 Pentose and glucuronate interconversions	0.90	0.53	0.00	0.01
39	map04146	过氧化物酶体 Peroxisome	1.36	0.88	0.00	0.01
28	map00592	α-亚麻酸代谢 alpha-Linolenic acid metabolism	0.97	0.60	0.00	0.02
20	map00360	苯丙氨酸代谢 Phenylalanine metabolism	0.69	0.40	0.00	0.03
24	map00460	氰胺酸代谢 Cyanoamino acid metabolism	0.83	0.50	0.00	0.03
16	map00906	类胡萝卜素生物合成 Carotenoid biosynthesis	0.56	0.31	0.00	0.05
16	map00950	异喹啉生物碱的生物合成 Isoquinoline alkaloid biosynthesis	0.56	0.31	0.00	0.04
23	map00130	泛醌和其他萜类醌的生物合 Ubiquinone and other terpenoid-quinone biosynthesis	0.80	0.50	0.00	0.05
32	map00260	甘氨酸、丝氨酸和苏氨酸代谢 Glycine, serine and threonine metabolism	1.11	0.78	0.00	0.06
23	map04712	植物昼夜节律 Circadian rhythm-plant	0.80	0.52	0.00	0.06
28	map00051	果糖和甘露糖代谢 Fructose and mannose metabolism	0.97	0.65	0.00	0.07
22	map00052	半乳糖代谢 Galactose metabolism	0.76	0.50	0.01	0.10
50	map00520	氨基糖和核苷酸糖代谢 Amino sugar and nucleotide sugar metabolism	1.74	1.34	0.01	0.10

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表 7 20个差异表达基因qRT-PCR的RQ值

Table 7. RQ values of qRT-PCR for 20 differentially expressed genes

基因 Gene	正常种 RQ值 RQ value in CK	平原种植退化种 RQ值 RQ value in BZ	调节 Regulation	RNA-seq调节 Regulation in RNA-seq
抗病反应蛋白 Disease resistance response protein	1.00 b	216.68 a	上调 Up	上调 Up
抗病蛋白 RGA2 Disease resistance protein RGA2	1.00 b	6.29 a	上调 Up	上调 Up
乙烯受体2 Ethylene receptor 2	1.00 a	0.48 b	下调 Down	下调 Down
过氧化氢酶1 Catalase 1	1.00 b	8.23 a	上调 Up	上调 Up
蔗糖合成酶 Sucrose synthase	1.00 b	10.50 a	上调 Up	上调 Up
抗病蛋白 RPS5 Disease resistance protein RPS5	1.00 b	10.02 a	上调 Up	上调 Up
抗病蛋白 Disease resistance protein	1.00 b	10.04 a	上调 Up	上调 Up
类烟草病毒增殖蛋白3 Tobamovirus multiplication protein 3-like	1.00 a	0.78 b	下调 Down	下调 Down
植物抗病反应蛋白 Plant disease resistance response protein	1.00 b	10.01 a	上调 Up	上调 Up
F-box家族蛋白 F-box family protein	1.00 b	7.815577 a	上调 Up	上调 Up
超氧化物歧化酶 [Cu-Zn] Superoxide dismutase [Cu-Zn]	1.00 b	9.217215 a	上调 Up	上调 Up
逆转录病毒相关 Pol多聚蛋白系-1 Retrovirus-related Pol polyprotein LINE-1	1.00 a	0.13 b	下调 Down	下调 Down
转座子 RE1的逆转录病毒相关 Pol多蛋白 Retrovirus-related Pol polyprotein from transposon RE1	1.00 a	0.01 b	下调 Down	下调 Down
转座子 TNT逆转录病毒相关 Pol多蛋白 Retrovirus-related Pol polyprotein from transposon TNT	1.00 a	0.06 b	下调 Down	下调 Down
BZIP转录因子家族蛋白 BZIP transcription factor family protein	1.00 a	0.46 b	下调 Down	下调 Down
过氧化物酶 Peroxidase	1.00 b	16.00 a	上调 Up	上调 Up
脱落酸受体 PYR1 Abscisic acid receptor PYR1	1.00 a	0.25 b	下调 Down	下调 Down
抗病蛋白 RPS2 Disease resistance protein RPS2	1.00 b	5.40 a	上调 Up	上调 Up
生长素反应因子1 Auxin response factor 1	1.00 a	0.50 b	下调 Down	下调 Down
烟草病毒增殖蛋白1 Tobamovirus multiplication protein 1	1.00 a	0.49 b	下调 Down	下调 Down
甘油醛-3-磷酸脱氢酶 GAPDH	1.00 a	1.00 a	-	-
注：同行数据后不同小写字母表示显著性差异（P<0.05）。 Note: Different lowercase letters in each line indicate significant difference at 0.05 level.

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