摘要: 目的Ⅲ类过氧化物酶（PRX）是一类植物特有的氧化还原酶，在植物生长发育和胁迫响应方面具有十分重要的作用。本研究拟探讨油橄榄Ⅲ类 PRX基因家族的进化和表达模式，旨在为油橄榄分子育种提供参考、�/sec> 方法利用生物信息学方法鉴定并分析了油橄榄Ⅲ类 PRX基因家族的系统发育关系、分组、染色体分布、复制基因、基序分布、基因结构图、顺式作用元件分布和在不同组织和不同非生物胁迫下的表达，并对部分成员基因进行了实时荧光定量PCR验证、�/sec> 结果�?）鉴定得到了106�?i>OePRX基因，根据其与拟南芥和毛果杨PRX蛋白序列的系统进化关系分为了14个组。（2（�i>OePRX基因不均匀地分布于23条染色体上；片段复制是该基因家族扩张的主要动力，且复制基因在进化过程中受到了较强的纯化选择；与拟南芥相比，油橄榃�i>PRX基因与毛果杨 PRX基因的亲缘关系更近。（3）同一组中OePRX蛋白的等电点、分子量、基序分布、基因结构、信号肽分布、在不同组织中的表达模式均比较保守； OePRX基因启动子中含有较多的生长发育元件和激素应答元件；在油橄榄干热胁迫和水涝胁迫下�?8%皃�i>OePRX基因显著差异表达、�/sec> 结论油橄榃�i>OePRX基因家族明显的进化扩张和多变的表达模式暗示了其功能的复杂性，尤其是复制基因的新功能化对油橄榄在地中海地区的广泛分布具有十分重要的适应性意义、�/sec>

关键诌�
• 油橄榃�/a> /
• Ⅲ类PRXs/
• 复制基因/
• 新功胼�/a> /
• 非生物胁�?/a>

Abstract: ObjectiveThe class � peroxidase is a group of plant-specific oxidoreductases, and plays an important role in plant development and stress response. This paper presents a detailed overview of evolution and expression of olive class � PRXgene family, aiming to provide a reference for olive molecular breeding in the future. MethodWith some bioinformatic tools, we finished the identification of the olive class � PRXgenes, completed the analyses of phylogenetic relationship, gene mapping on the chromosomes, pairs of duplicates, motifs, gene structure, cis-acting elements and gene expression in different tissues or biotic stresses, and did a verification of RNA-seq by RT-qPCR. Result(1) 106 OePRXgenes were obtained, and classified into 14 groups based on the phylogeny with AtPRX and PtPRX. (2) OePRXgenes were unevenly located on 23 chromosomes, and ragment replication was the main driving force for the expansion of gene family. Compared with AtPRX, OePRXhad a closer relationship with PtPRX. (3) The characterization of pI, MW, motifs, gene structure, signal peptide, and expression in different tissues had the expected group-conserved patterns. The promoters of OePRXcontained a variety of development and harmone elements; 38% of OePRXgenes was differentially expressed in heat, drought and waterlogged stress. ConclusionThe differential expansion and differential expression patterns may imply flexibility in neofunctionalization of duplicated class � peroxidase genes, which is of adaptive significance to the strong resistance of olive to a diversity of conditions, hence contributing to the importance of olive as a Mediterranean Basin staple food.

Key words:
• Olea europaea/
• class � peroxidase/
• replicator/
• new function/
• abiotic stress

�?nbsp; 1油橄榄、毛果杨和拟南芥Ⅲ类PRX蛋白的系统进化关糺�/p>

Figure 1.Phylogeny of class � PRX proteins from olive, black cottonwood andArabidopsis thaliana

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�?nbsp; 2Ⅲ类油橄榃�i>PRX基因在染色体上的分布国�/p>

串联复制基因簇用方框标记。Tandem replication gene cluster is marked with a box.

Figure 2.Chromosomal distribution ofPRXgenes in class � olive

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�?nbsp; 3油橄榄Ⅲ籺�i>PRX的共线性分枏�/p>

Figure 3.Collinearity analysis of olive type �PRXgenes

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�?nbsp; 4油橄榄、毛果杨和拟南芥Ⅲ类PRX的共线性分枏�/p>

Figure 4.Collinearity analysis of class �PRXin olive, black cottonwood andArabidopsis thaliana

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�?nbsp; 5油橄榄Ⅲ类PRX各复制基因对的Ka、Ks分析

Ka. 非同义替换；Ks. 同义替换；Oe-At. 油橄榄和拟南芥之间的PRX复制基因对；Oe-Pt. 油橄榄和毛果杨之间的PRX复制基因对。Ka, nonsynonymous substitution rate; Ks, synonymous substitution rate; Oe-At,PRXduplicate gene pairs between olive andArabidopsis thaliana; Oe-Pt,PRXduplicate gene pairs between olive and black cottonwood.

Figure 5.Ka, Ks analyses of olive class �PRXduplicated genes

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�?nbsp; 6油橄榄Ⅲ籺�i>PRX基因的系统进化树（A）、保守基序（B）和基因结构图（C（�/p>

Figure 6.Phylogeny (A), conserved motif (B) and extron-intron structure (C) of the olive class �PRXgenes

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�?nbsp; 7油橄榄Ⅲ籺�i>PRX的启动子上的顺式作用元件分析

Figure 7.cis-element analysis of promoters of olive class �PRXgenes

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�?nbsp; 8油橄榄Ⅲ籺�i>PRX基因在不同器官或组织中的表达模式

模块A. 在不同组织中均高表达；模块B. 在不同组织中均有较高表达；模块C. 在不同组织中均较低表达；模块D. 在花中特异高表达；模块E. 在除果外的组织中高表达；模块F. 在除果外的组织较高表�? 模块G. 在根中特异高表达；模块H. 在各组织中低表达或几乎不表达。Module A, high expression in all tissues; module B, relative high expression in all tissues; module C, relative low expression in all tissues; module D, high specific expression in flower; module E, high expression in all tissues except fruit; module F, relative high expression in all tissues except fruit; module G, high specific expression in root; module H, low or no expression in all tissues.

Figure 8.Expression patterns of olive class �PRXgenes in different organs or tissues

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�?nbsp; 9RT-qPCR验证转录组数�?/p>

柱状图为运用RNA-seq得到的tpm值（相对表达�?）。折线图为运用RT-qPCR得到�?^−ΔΔCT值（相对表达�?）。Bars indicate tpm value measured by RNA-seq (relative expression 1). Lines indicate 2^−ΔΔCTmeasured by RT-qPCR (relative expression 2).

Figure 9.RT-qPCR validations of RNA-seq data

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�?nbsp; 10油橄榄Ⅲ籺�i>PRX基因响应干热和水涝胁迫的差异表达模式

Day（−3�? 干旱处理�?天；Day13. 干旱处理�?3天且伴随高温；Day27. 干旱处理�?7天；Day80. 复水�?2天。模�?. 干旱期间基因表达下调，复水后上调；模�?. 干旱期间基因表达上调，复水后下调；模�?. 后期干旱期间基因表达上调，后期干旱及复水后下调；模块4. 仅在高温下基因表达上调，高温胁迫后，基因表达下调。复制基因名称的颜色与系统进化分组的颜色相对应。Day（−3�? 3 d before drought treatment; Day13, 13th day of drought treatment with high temperature; Day27, 27th day after drought treatment; Day80, 52 d after re-irrigation. Module 1, gene expression is down regulated during drought and up regulated after rehydration; module 2, gene expression is upregulated during drought and downregulated after rehydration; module 3, gene expression is upregulated during late drought, and downregulated after late drought and rehydration; module 4, gene expression is upregulated only under high temperature, and downregulated after high temperature stress. The color of the replicated gene name corresponds to the color of the phylogenetic grouping.

Figure 10.Differential expression patterns of class �PRXgenes in olive oil in response to dry heat and waterlogging stresses

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�?nbsp; 1油橄榄Ⅲ籺�i>PRX基因的命名及其理化性质

Table 1.Nomenclature and physicochemical properties of class �PRXgenes from olive

名称 Name	ID	内含子相� Intron phase	蛋白质长� Protein length/aa	分子� Molecular mass/kDa	等电� Isoelectric point	信号� Signal peptide	预测的亚细胞定位 Predicted subcellular localization
OePRX1	EVM0022477	0	344	38.40	6.10	� No	胞外 Extracellular
OePRX2	EVM0032926	0	266	29.38	8.97	� No	细胞� Cytosol
OePRX3	EVM0020804	0	327	35.65	9.21	� Yes	叶绿� Chloroplast
OePRX4	EVM0047797	100	327	35.92	8.24	� No	细胞� Nucleus
OePRX5	EVM0009292	100	316	34.40	8.72	� Yes	液泡� Vacular membrane
OePRX6	EVM0060238	0	330	35.97	8.89	� No	叶绿� Chloroplast
OePRX7	EVM0022620	12001000	276	30.39	8.66	� No	细胞� Cytosol
OePRX8	EVM0017319	1200201	279	30.58	7.70	� No	细胞� Cytosol
OePRX9	EVM0047458	20010	250	27.43	5.67	� No	细胞� Cytosol
OePRX10	EVM0057886	0	319	34.48	6.98	� Yes	叶绿� Chloroplast
OePRX11	EVM0047784	0	339	37.00	7.48	� Yes	质膜 Plasma membrane
OePRX12	EVM0039807	20	358	38.84	7.51	� Yes	叶绿� Chloroplast
OePRX13	EVM0011450	0	317	34.35	9.14	� Yes	叶绿� Chloroplast
OePRX14	EVM0019223	1	309	35.27	9.41	� Yes	胞外 Extracellular
OePRX15	EVM0012728	1	320	36.91	9.20	� Yes	细胞� Cytosol
OePRX16	EVM0000231	0	336	37.28	7.08	� Yes	叶绿� Chloroplast
OePRX17	EVM0024793	0	322	35.21	8.38	� Yes	叶绿� Chloroplast
OePRX18	EVM0042519	0	321	34.96	8.76	� Yes	叶绿� Chloroplast
OePRX19	EVM0059982	0	321	34.90	8.77	� Yes	叶绿� Chloroplast
OePRX20	EVM0019685	0	333	36.73	5.68	� No	质膜 Plasma membrane
OePRX21	EVM0058087	0	324	35.74	8.53	� Yes	细胞� Cytosol
OePRX22	EVM0027348	0	323	35.02	5.82	� Yes	液液泡膜 Vacular membrane
OePRX23	EVM0060830	0	331	36.34	9.41	� Yes	胞外 Extracellular
OePRX24	EVM0015621	0	247	27.39	8.88	� No	细胞� Nucleus
OePRX25	EVM0010445	12101221	327	35.56	9.26	� No	叶绿� Chloroplast
OePRX26	EVM0006524	0	338	37.53	6.45	� Yes	胞外 Extracellular
OePRX27	EVM0042768	22	304	32.63	6.58	� Yes	叶绿� Chloroplast
OePRX28	EVM0048359	0	355	37.59	8.35	� Yes	叶绿� Chloroplast
OePRX29	EVM0028664	0	317	34.49	9.09	� Yes	液泡� Vacular membrane
OePRX30	EVM0035621	0	327	34.85	5.51	� Yes	胞外 Extracellular
OePRX31	EVM0048816	100	324	36.67	6.26	� Yes	胞外 Extracellular
OePRX32	EVM0003637	1002	350	39.07	8.83	� Yes	胞外 Extracellular
OePRX33	EVM0019330	0	333	36.44	9.48	� Yes	液泡� Vacular membrane
OePRX34	EVM0033281	2100102	273	30.15	7.73	� No	细胞� Cytosol
OePRX35	EVM0015060	10100000	319	35.73	6.63	� No	叶绿� Chloroplast
OePRX36	EVM0017946	0	329	37.29	8.48	� Yes	细胞� Cytosol
OePRX37	EVM0021366	∑�/td>	331	36.79	9.34	� Yes	叶绿� Chloroplast
OePRX38	EVM0060527	0	342	37.64	6.21	� Yes	细胞� Cytosol
OePRX39	EVM0014494	0	321	35.37	6.18	� Yes	胞外 Extracellular
OePRX40	EVM0042196	0	319	34.12	9.41	� Yes	叶绿� Chloroplast
OePRX41	EVM0015681	0	346	38.94	8.86	� Yes	内质� Endoplasmic reticulum
OePRX42	EVM0047610	0	327	35.27	8.80	� Yes	叶绿� Chloroplast
OePRX43	EVM0047452	20002	379	41.95	8.87	� No	叶绿� Chloroplast
OePRX44	EVM0061929	0	319	34.23	4.45	� Yes	胞外 Extracellular
OePRX45	EVM0054999	0	291	32.27	5.44	� No	胞外 Extracellular
OePRX46	EVM0048606	0	328	35.67	6.51	� Yes	液泡� Vacular membrane
OePRX47	EVM0037058	20	315	34.41	7.55	� No	内质� Endoplasmic reticulum
OePRX48	EVM0022847	20	413	45.85	9.56	� No	叶绿� Chloroplast
OePRX49	EVM0020385	20	373	41.16	9.35	� No	叶绿� Chloroplast
OePRX50	EVM0045080	0	317	34.55	9.02	� Yes	叶绿� Chloroplast
OePRX51	EVM0044622	0	320	35.53	9.54	� Yes	叶绿� Chloroplast
OePRX52	EVM0021425	0	324	36.14	6.17	� Yes	胞外 Extracellular
OePRX53	EVM0045494	0	327	35.52	8.71	� Yes	液泡� Vacular membrane
OePRX54	EVM0020915	20	306	33.46	8.90	� Yes	液泡� Vacular membrane
OePRX55	EVM0031491	0	314	34.07	9.32	� Yes	叶绿� Chloroplast
OePRX56	EVM0005944	2	292	31.79	9.45	� Yes	叶绿� Chloroplast
OePRX57	EVM0015088	0	317	34.42	9.13	� Yes	胞外 Extracellular
OePRX58	EVM0050632	0	314	34.04	9.25	� Yes	叶绿� Chloroplast
OePRX59	EVM0028088	0	317	34.23	8.99	� Yes	胞外 Extracellular
OePRX60	EVM0025336	0	326	34.42	7.58	� No	叶绿� Chloroplast
OePRX61	EVM0012469	0	317	34.68	9.33	� Yes	胞外 Extracellular
OePRX62	EVM0049574	0	340	37.08	6.25	� Yes	胞外 Extracellular
OePRX63	EVM0050273	10	240	26.55	6.22	� No	叶绿� Chloroplast
OePRX64	EVM0050817	0	310	34.15	8.46	� Yes	叶绿� Chloroplast
OePRX65	EVM0009492	0	322	34.85	4.99	� Yes	叶绿� Chloroplast
OePRX66	EVM0047271	1200201	333	36.96	8.64	� No	质膜 Plasma membrane
OePRX67	EVM0005889	210122	509	58.00	8.73	� No	细胞� Nucleus
OePRX68	EVM0026529	0	331	36.43	9.50	� Yes	液泡� Vacular membrane
OePRX69	EVM0038965	20	310	33.90	9.23	� Yes	质膜 Plasma membrane
OePRX70	EVM0024841	0	328	35.72	8.37	� Yes	叶绿� Chloroplast
OePRX71	EVM0047959	3	314	34.07	6.94	� Yes	叶绿� Chloroplast
OePRX72	EVM0059042	0	339	37.43	5.56	� Yes	质膜 Plasma membrane
OePRX73	EVM0022158	0	318	34.18	6.88	� Yes	叶绿� Chloroplast
OePRX74	EVM0027175	0	326	36.14	9.75	� Yes	叶绿� Chloroplast
OePRX75	EVM0057531	20	287	30.73	8.89	� Yes	液泡� Vacular membrane
OePRX76	EVM0032201	0	293	32.34	8.75	� No	叶绿� Chloroplast
OePRX77	EVM0059919	0	339	37.61	8.36	� Yes	液泡� Vacular membrane
OePRX78	EVM0011789	0	355	39.51	8.06	� No	质膜 Plasma membrane
OePRX79	EVM0024970	∑�/td>	329	36.54	9.19	� Yes	叶绿� Chloroplast
OePRX80	EVM0047604	100	339	37.82	8.30	� No	细胞� Cytosol
OePRX81	EVM0061300	0	330	36.94	5.94	� Yes	叶绿� Chloroplast
OePRX82	EVM0042525	0	298	32.64	8.62	� Yes	胞外 Extracellular
OePRX83	EVM0045066	0	318	34.03	9.15	� Yes	叶绿� Chloroplast
OePRX84	EVM0029808	0	354	39.38	5.68	� No	胞外 Extracellular
OePRX85	EVM0032932	1022010	247	27.30	5.65	� No	细胞� Cytosol
OePRX86	EVM0038819	2001020	250	27.72	5.44	� No	细胞� Cytosol
OePRX87	EVM0002745	200	321	34.58	4.77	� No	细胞� Nucleus
OePRX88	EVM0038873	200	326	34.71	8.30	� Yes	液泡� Vacular membrane
OePRX89	EVM0026260	2	348	38.22	9.56	� Yes	质膜 Plasma membrane
OePRX90	EVM0044542	0	296	32.61	8.14	� Yes	胞外 Extracellular
OePRX91	EVM0046395	0	318	34.24	9.54	� Yes	叶绿� Chloroplast
OePRX92	EVM0042595	0	316	33.95	7.51	� Yes	叶绿� Chloroplast
OePRX93	EVM0047351	0	320	35.08	5.48	� Yes	叶绿� Chloroplast
OePRX94	EVM0045920	0	332	35.75	4.63	� Yes	胞外 Extracellular
OePRX95	EVM0044144	0	325	35.63	8.75	� No	叶绿� Chloroplast
OePRX96	EVM0048820	0	314	35.88	8.61	� Yes	细胞� Cytosol
OePRX97	EVM0058650	0	350	38.67	6.94	� Yes	叶绿� Chloroplast
OePRX98	EVM0026875	0	371	40.68	6.31	� No	叶绿� Chloroplast
OePRX99	EVM0019820	0	340	36.88	6.59	� Yes	胞外 Extracellular
OePRX100	EVM0018143	20020200000	394	43.59	9.28	� No	叶绿� Chloroplast
OePRX101	EVM0029423	0	343	37.01	8.94	� Yes	叶绿� Chloroplast
OePRX102	EVM0036633	0	342	37.59	9.17	� Yes	胞外 Extracellular
OePRX103	EVM0041887	1	312	33.33	5.97	� Yes	叶绿� Chloroplast
OePRX104	EVM0032297	0	327	35.71	8.57	� Yes	胞外 Extracellular
OePRX105	EVM0042746	0	302	33.53	8.96	� Yes	胞外 Extracellular
OePRX106	EVM0051394	0	343	37.01	8.94	� Yes	叶绿� Chloroplast

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�?nbsp; 26个油橄榄Ⅲ类PRX基因RT-qPCR的引�?/p>

Table 2.Primers of six olive class �PRXgenes used for RT-qPCR

基因名称 Gene name	引物�?′�?′） Primer sequence (5′−3�?	基因产物长度 Gene product length/bp
OePRX1	F: TCCAGGAGTTGTTTCTTGTGC	114
	R: CCTTCTTCCGTCTTTTCTTCC
OePRX8	F: AAGGGCGCATCCTGAAAG	200
	R: AGAAGGCATCTTCATCCTTAGC
OePRX33	F: GGAAGCATAATAAGCGAGAAG	235
	R: TTATTGGAGCCACTCAAACTG
OePRX36	F: ACAAACGCCACAAGAACACTG	195
	R: CACTCCCTCTCTAAAGCCTCC
OePRX66	F: GATTCGCAATGAGGAGGAGTA	140
	R: AGCAACAACACCAGCAAGC
OePRX99	F: GCTCGGGCTTTTAGAATCATC	204
	R: GAAGGAAGGTTTGCCAGTGTT

下载: 导出CSV

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