摘要: 目的为研究蔷薇属植物响应黑斑病侵染的稳定表达的最适内参基因，解析茉莉酸在蔷薇属植物响应蔷薇盘二孢侵染过程中的作用、�/sec> 方法以黑斑病病原菌蔷薇盘二孢侵染�?个蔷薇属�?品种不同时间的离体叶片为材料，利用qRT-PCR技术及geNorm、NormFinder、BestKeeper软件�?个候选内参基因（ ACT�?i>GAPDH�?i>PP2 A�?i>Rcl2�?i>SAND�?i>TIP�?i>TUA�?i>TUB�?i>UBC）的表达量进行测定和分析，利用筛选出的内参基因，对蔷薇属植物茉莉酸（JA）抗病途径相关基因＇�i>COI1�?i>OPR3�?i>MYC2�?i>JAR1）的表达水平进行定量分析、�/sec> 结果�?（�i>UBC可作�?种蔷薇属植物共同适用的内参基因，可用于后续分析JA抗病途径相关基因的表达水平。（2）内源JA含量�?种蔷薇属植物响应蔷薇盘二孢侵染的过程中存在差异。在黑斑病高抗植物受侵染0 ~ 4 d间JA含量下调�? ~ 8 d间上调。在黑斑病易感植物受侵染0 ~ 8 d间，内源JA含量呈下调趋势。（3）JA合成相关基因 OPR3叉�i>JAR1表达量在受侵染初期表达量趋势存在差异。在除荷花蔷薇外的其�?种植物中＋�i>OPR3在侵染初期（0 ~ 0.5 d）表达下调， JAR1在侵染初期表达上调、�i>OPR3咋�i>JAR1在侵染后期均表达上调，黑斑病易感材料的上调程度高于高抗材料。（4）JA信号传导相关基因 COI1叉�i>MYC2在受侵染初期表达量趋势同样存在差异、�i>COI1在黑斑病高抗材料受侵染初期上调表达，在黑斑病易感材料下调表达＋�i>MYC2�?种植物受侵染0 ~ 2 d中均下调表达、�i>COI1叉�i>MYC2表达量在受侵�? d后均上调表达，且在黑斑病易感植物中的上调程度大于黑斑病高抗材料、�/sec> 结论与JA信号传导相关皃�i>MYC2�?i>COI1在蔷薇属植物抵御黑斑病病菌入侵过程中发挥负调控作用，且由JA通路介导的抵御死体营养型病原菌的侵染在后期发挥了作用、�/sec>

关键诌�
• 蔷薇屝�/a> /
• 蔷薇盘二孡�/a> /
• 内参基因筛逈�/a> /
• 茉莉酷�/a> /
• 表达分析

Abstract: ObjectiveThis paper aims to select the appropriate reference gene and analyze the role of jasmonic acid in Rosaspecies and cultivars responding to Marssonina rosae. Method R. multifloraf. carnea, R. xanthinaf. spontanea, R. glauca, R. rugosa, R. hybrida‘Porcelina� and R12-26 (hybrid of R. rugosaand R. hybrida‘Porcelina�? were used as materials, qRT-PCR technology, geNorm, NormFinder, BestKeeper were used to evaluate the expression stability of nine candidate reference genes ( ACT, GAPDH, PP2 A, RCl2, SAND, TIP, TUA, TUBand UBC). The expression levels of jasmonic acid resistance pathway relatedgenes ( COI1, OPR3, MYC2 and JAR1) in six Rosaspecies and cultivars were quantitatively analyzed. Result(1) UBCwas the common reference gene of six Rosaspecies and cultivars responding to M. rosae, and can be used for analyzing the related gene expression levels to JA resistance pathway. (2) There were differences in JA content among the six plants responding to M. rosae. In disease-resistant plants, JA concentration decreased at 0�? d and increased at 4�? d. In disease-susceptible plants, JA concentration decreased at 0�? d and increased significantly at 8�?0 d. (3) The expression levels of JA synthesis-related genes OPR3 and JAR1 were different at the early stage of infection. The expression level of OPR3 was down-regulated and JAR1 up-regulated in five plants except for R. multifloraf. carnea. OPR3 and JAR1 were all up-regulated in the late stage of infection, however the up-regulation degree in black-spot susceptible species was significantly higher than that of resistant species. (4) There were also differences in the expression levels of JA signaling related genes COI1 and MYC2 at the early stage of infection. The expression of COI1 was up-regulated in disease-resistant plants and down-regulated in disease-susceptible plants at the early stage of infection, MYC2 was down-regulated in 6 plants infected 0�? d. COI1 and MYC2 were up-regulated after 2 d of infection, and the up-regulation degree in black-spot susceptible species was significantly higher than that of resistant species. Conclusion MYC2 and COI1, which are related to JA signaling, play a negative regulatory role in the resistance of Rosaplants to the invasion of black spot pathogens, and the JA pathway mediated resistance to the invasion of dead vegetative pathogens plays a role in the later stage of infection.

Key words:
• Rosa/
• Marssonina rosae/
• reference gene selection/
• jasmonic acid/
• expression analysis

�?nbsp; 16种蔷薇属植物离体叶片侵染蔷薇盘二孢的发病情况

红色圆圈表示病斑开始出现。黑色圆圈表示病斑蔓延至整个叶片。Red circle indicates the disease onset with the appearance brown necrotic lesions. Black circles indicate the spread of the brown necrotic lesions throughout the whole leaf.

Figure 1.Incidence of in vitro leaves infected byM. rosaeof 6Rosaspecies and cultivars

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�?nbsp; 29个候选内参基因响应蔷薇盘二孢侵染时的Ct倻�/p>

Figure 2.Ct value of 9 candidate reference genes in response toM. rosae

下载: 全尺寸图牆�/a> 幻灯牆�/a>

�?nbsp; 3利用geNorm软件计算的候选内参基因的表达稳定性及配对变异倻�/p>

M代表各候选内参基因的表达稳定度。下同、�i>Mrepresents the average expression stability of each candidate reference gene. The same below.

Figure 3.Average expression stability values and variation values of candidate reference genes calculated by geNorm

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�?nbsp; 4Normfinder软件计算获得的候选内参基因的表达稳定�?/p>

Figure 4.Expression stability of candidate reference genes and by pairwise variation using Normfinder

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�?nbsp; 5BestKeeper软件计算获得的候选内参基因的表达稳定�?/p>

SD代表各候选内参基因Ct值的标准偏差。SD represents the standard deviation of Ct value of each candidate reference gene.

Figure 5.Expression stability of candidate reference genes calculated by BestKeeper

下载: 全尺寸图牆�/a> 幻灯牆�/a>

�?nbsp; 6内源茉莉酸在蔷薇属植物响应蔷薇盘二孢侵染的含野�/p>

Figure 6.Content of JA inRosaspecies and cultivars responding toM. rosae

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�?nbsp; 7茉莉酸相关基因在蔷薇盘二孢侵染不同时间的相对表达野�/p>

Figure 7.Relative expression level of JA-related genes at different time ofM. rosaeinfection

下载: 全尺寸图牆�/a> 幻灯牆�/a>

�?nbsp; 1基因引物基本信息

Table 1.Basic information of gene primers

基因 Gene	引物序列�?′�?′） Primer sequence (5�?3�?	退火温�?br/>Annealing temperature (Tm)/ℂ�/td>	扩增效率 Amplification efficiency (E)/%	r
ACT	F: TCAAGGATTGGTGGACTTCAGT R: ACCAGAGAACAAGAATGCAAGC	55	96.0	0.970
GAPDH	F: TATGACCAGATCAAGGCTGCT R: ACCAATGAAGTCGGTTGACAC	55	102.5	0.999
PP2A	F: TGTCACTGCATCAAAGGACAG R: GACGAATTGTCTTCTCCACCA	55	102.0	0.982
Rcl2	F: ATGGGAAATGCCCTACCT R: CACTTGTCCGACTGTTGC	55	95.7	0.969
SAND	F: BGTGTTGAGGAGTTGCCTCTTG R: AACCTGTCGGGAGAATCTGTT	55	98.5	0.989
TIP	F: GAATCCACGGCTGGGAAA R: CAGTTCGTGGGTGGAGGAGTT	55	104.2	0.998
TUA	F: CATTGAGCGTCCCACCTA R: CACATCCACATTCAGAGCC	55	103.2	0.987
TUB	F: GTACATGGCCTGCTGTTTGAT R: ATGGTACGCTTGGTCTTGATG	55	96.1	0.972
UBC	F: GCCAGAGATTGCCCATATGTA R: TCACAGAGTCCTAGCAGCACA	55	103.5	0.996
COI1	F: AATGAGGGGCTGTTGCTTCA R: GATCCCCTGTACCCTTGCAC	56	102.2	0.984
MYC2	F: CGGCAGCAGCGTCAAGAAT R: GAGGTCGGAGTGGTGGGAAT	57	101.0	0.957
OPR3	F: CATCAGCGAAGGCACTTTGG R: GGCGTCGACTACCTTCTTCC	55	103.6	0.951
JAR1	F: TTGGGTGCTACTTCTTTCTCAG R: AATTGGAGGAAGGAGGGTG	57	104.4	0.952

下载: 导出CSV

�?nbsp; 29个候选内参基因的表达稳定性排庎�/p>

Table 2.Expression stability ranking of 9 candidate internal reference genes

�?品种 Species/cultivar	方法 Method	1	2	3	4	5	6	7	8	9
荷花蔷薇 R. multifloraf.carnea	geNorm	UBC	TIP	PP2A	GAPDH	TUB	ACT	SAND	TUA	Rcl2
	Normfinder	UBC	TIP	PP2A	GAPDH	TUB	ACT	TUA	SAND	Rcl2
	Bestkeeper	TIP	ACT	UBC	TUB	SAND	PP2A	TUA	GAPDH	Rcl2
	RefFinder	UBC	TIP	PP2A	TUB	ACT	GAPDH	SAND	TUA	Rcl2
单瓣黄刺�?br/>R. xanthinaf.spontanea	geNorm	UBC	PP2A	SAND	TIP	GAPDH	ACT	Rcl2	TUB	TUA
	Normfinder	PP2A	UBC	TIP	SAND	GAPDH	ACT	Rcl2	TUB	TUA
	Bestkeeper	PP2A	TIP	UBC	ACT	GAPDH	SAND	Rcl2	TUB	TUA
	RefFinder	PP2A	UBC	TIP	SAND	GAPDH	ACT	Rcl2	TUB	TUA
玫瑰 R. rugosa	geNorm	UBC	TIP	GAPDH	TUB	TUA	PP2A	SAND	Rcl2	ACT
	Normfinder	UBC	TIP	PP2A	GAPDH	TUB	SAND	TUA	Rcl2	ACT
	Bestkeeper	UBC	TIP	TUA	TUB	GAPDH	SAND	ACT	PP2A	Rcl2
	RefFinder	UBC	TIP	GAPDH	TUB	TUA	PP2A	SAND	ACT	Rcl2
R12-26	geNorm	PP2A	TIP	UBC	SAND	GAPDH	TUB	TUA	ACT	Rcl2
	Normfinder	TIP	GAPDH	UBC	PP2A	SAND	TUB	TUA	ACT	Rcl2
	Bestkeeper	UBC	ACT	GAPDH	TIP	PP2A	SAND	Rcl2	TUB	TUA
	RefFinder	UBC	TIP	PP2A	GAPDH	SAND	ACT	TUB	TUA	Rcl2
红叶蔷薇 R. glauca	geNorm	UBC	TIP	SAND	PP2A	TUA	TUB	GAPDH	Rcl2	ACT
	Normfinder	UBC	TIP	TUA	SAND	PP2A	TUB	GAPDH	Rcl2	ACT
	Bestkeeper	SAND	TIP	PP2A	UBC	ACT	TUA	TUB	GAPDH	Rcl2
	RefFinder	UBC	TIP	SAND	PP2A	TUA	TUB	GAPDH	ACT	Rcl2
‘波塞妮娜‘�br/>R. hybrida‘Porcelina‘�/td>	geNorm	SAND	TIP	UBC	GAPDH	PP2A	ACT	TUB	Rcl2	TUA
	Normfinder	GAPDH	TIP	UBC	SAND	TUB	PP2A	ACT	Rcl2	TUA
	Bestkeeper	PP2A	UBC	SAND	TIP	GAPDH	ACT	TUB	Rcl2	TUA
	RefFinder	UBC	TIP	SAND	GAPDH	PP2A	TUB	ACT	Rcl2	TUA

下载: 导出CSV

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