苹果砧木‘青砧1号’叶片cDNA文库构建及MdMLO上游调控因子的筛选

doi:10.16420/j.issn.0513-353x.2022-0159

摘要/Abstract

摘要：

MLO（Mildew Resistance Locus O）家族基因是植物特有的“感病基因”，普遍认为它对白粉病具有负调控作用。目前在单子叶和双子叶植物中发现许多抗白粉病MLO基因，但对于其上游调控因子如何调控其表达而影响白粉病发生的研究却很少。本研究中以易感白粉病的苹果砧木‘青砧1号’叶片为材料，构建其cDNA文库，文库滴度4 × 10⁹ cfu · mL^-1，重组率为100%。生物信息分析苹果MdMLO家族基因启动子区共有38种顺式作用元件，包含激素、应激防御和生长发育等响应元件。以TCA和TC-rich repeats为诱饵序列构建诱饵载体，分别命名为Bait-TCA和Bait-TC。利用酵母单杂交的方法筛选其上游调控因子，结果显示Bait-TCA具有自激活现象无法筛选，Bait-TC诱饵载体调取获得30条基因序列，其中6条参与植物抗逆防御反应，推测它们可能通过与TC-rich repeats元件互作来调控MLO的表达，进而调控苹果叶片对于白粉病的响应。

关键词: 苹果, MLO基因, cDNA文库, 酵母单杂交, 顺式作用元件

Abstract:

MLO is a plant specific recessive resistance gene family. It is widely believed that it has a negative regulatory effect on powdery mildew. Many powdery mildew resistance MLO genes have been found in both monocotyledon and dicotyledon. However，there are few studies on the mechanism of upstream regulator of MLO genes. In this study，a cDNA library of apple rootstock‘Qingzhen 1’leaf was constructed with a titer of about 4 × 10⁹ cfu · mL^-1，and the recombination rate was 100%. Bioinformatics analysis of apple MLO gene family promoter region showed that there were 38 kinds of cis-acting elements，including hormone，stress defense response as well as growth and development response elements，etc. TCA and TC-rich repeats cis-acting elements were selected as bait sequences and constructed bait vectors named Bait-TCA and Bait-TC for yeast one-hybrid，respectively. The results showed that the Bait-TCA had self-activation and could not be used for screening. Among the Bait-TC obtained 30 sequences，six sequences were involved in plant anti-stress defense response. It suggested that they might be involved in the response of apple to powdery mildew through regulating the expression of MLO genes.

Key words: apple, MLO gene, cDNA library, yeast one-hybrid, cis-acting element

中图分类号:

S666.1

张琨, 思彬彬, 周军, 任玉锋, 张欣, 徐文娣, 王佳伟, 乔帅, 王惠冉. 苹果砧木‘青砧1号’叶片cDNA文库构建及MdMLO上游调控因子的筛选[J]. 园艺学报, 2023, 50(5): 933-946.

ZHANG Kun, SI Binbin, ZHOU Jun, REN Yufeng, ZHANG Xin, XU Wendi, WANG Jiawei, QIAO Shuai, WANG Huiran. Construction of cDNA Library of Apple Rootstock‘Qingzhen 1’Leaf and Screen of MdMLO Genes’ Upstream Regulator[J]. Acta Horticulturae Sinica, 2023, 50(5): 933-946.

图/表 11

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元件名称 Element name	元件序列 Element sequence	功能 Function
AAAC-motif	CAATCAAAACCT	光响应元件 Light responsive element
ACE	GACACGTATG	参与光反应元件 cis-Acting element involved in light responsiveness
AE-box	AGAAACAA	光响应组件的一部分 Part of a module for light response
AT1-motif	AATTATTTTTTATT	光响应组件的一部分 Part of a light responsive module
ATCT-motif	AATCTAATCC	光响应DNA保守元件的一部分 Part of a conserved DNA module involved in light responsiveness
chs-Unit 1 m1	ACCTAACCCGG	光响应元件的一部分 Part of a light responsive element
G-Box	CACGTT	光反应元件 cis-Acting regulatory element involved in light responsiveness
MRE	AACCTAA	MYB结合位点参与光响应 MYB binding site involved in light responsiveness
GARE-motif	TCTGTTG	赤霉素响应元件 Gibberellin-responsive element
AuxRE	TGTCTCAATAAG	生长素响应元件的一部分 Part of an auxin-responsive element
AuxRR-core	GGTCCAT	生长素响应元件 cis-Acting regulatory element involved in auxin responsiveness
ABRE	ACGTG	脱落酸响应元件 cis-Acting element involved in the abscisic acid responsiveness
P-box	CCTTTTG	赤霉素响应元件 Gibberellin-responsive element
TGACG-motif	TGACG	茉莉酸甲酯响应元件 cis-Acting regulatory element involved in the MeJA-responsiveness
TGA-element	AACGAC	生长素响应元件 Auxin-responsive element
TATC-box	TATCCCA	赤霉素响应元件 cis-Acting element involved in gibberellin-responsiveness
circadian	CAAAGATATC	响应昼夜节律控制元件 cis-Acting regulatory element involved in circadian control
AACA_motif	TAACAAACTCCA	胚乳特异性负表达元件 Involved in endosperm-specific negative expression
AT-rich element	ATAGAAATCAA	AT富集DNA结合蛋白位点（ATBP-1） Binding site of AT-rich DNA binding protein（ATBP-1）
CAT-box	GCCACT	分生组织表达相关元件 cis-Acting regulatory element related to meristem expression
GCN4_motif	TGAGTCA	胚乳表达响应元件 cis-Regulatory element involved in endosperm expression
HD-Zip 1	CAAT(A/T)ATTG	栅栏叶肉细胞分化响应元件 Element involved in differentiation of the palisade mesophyll cells
HD-Zip 3	GTAAT(G/C)ATTAC	蛋白结合位点 Protein binding site
MSA-like	TCCAACGGT	细胞周期调节元件cis-Acting element involved in cell cycle regulation
O2-site	GATGATGTGG	玉米醇溶蛋白代谢调节元件cis-Acting regulatory element involved in zein metabolism regulation
RY-element	CATGCATG	种子特异调控元件 cis-Acting regulatory element involved in seed-specific regulation
Unnamed_1	GAATTTAATTAA	60K蛋白结合位点 60K protein binding site
DRE	TACCGACAT	脱水、低温、盐响应元件 cis-Acting element involved in dehydration，low-temp，salt stresses
CCAAT-box	CAACGG	HvMYB1结合位点 HvMYB1 binding site
GC-motif	CCCCCG	增强缺氧特异性诱导元件 Enhancer-like element involved in anoxic specific inducibility
LTR	CCGAAA	低温响应元件 cis-Acting element involved in low-temperature responsiveness
MBS	CAACTG	MYB结合位点参与干旱诱导 MYB binding site involved in drought-inducibility
TCA-element	CCATCTTTTT	水杨酸响应元件 cis-Acting element involved in salicylic acid responsiveness
TC-rich repeats	GTTTTCTTAC	响应防御和应激反应元件 cis-Acting element involved in defense and stress responsiveness
ARE	AAACCA	厌氧诱导关键元件 cis-Acting regulatory element essential for the anaerobic induction
WUN-motif	AAATTTCCT	干旱响应元件Wound-responsive element
CAAT-box	CCAAT	启动子增强子区域元件Common cis-acting element in promoter and enhancer regions
TATA-box	TATA	启动子核心元件Core promoter element around -30 of transcription start

元件名称 Element name	元件序列 Element sequence	功能 Function
AAAC-motif	CAATCAAAACCT	光响应元件 Light responsive element
ACE	GACACGTATG	参与光反应元件 cis-Acting element involved in light responsiveness
AE-box	AGAAACAA	光响应组件的一部分 Part of a module for light response
AT1-motif	AATTATTTTTTATT	光响应组件的一部分 Part of a light responsive module
ATCT-motif	AATCTAATCC	光响应DNA保守元件的一部分 Part of a conserved DNA module involved in light responsiveness
chs-Unit 1 m1	ACCTAACCCGG	光响应元件的一部分 Part of a light responsive element
G-Box	CACGTT	光反应元件 cis-Acting regulatory element involved in light responsiveness
MRE	AACCTAA	MYB结合位点参与光响应 MYB binding site involved in light responsiveness
GARE-motif	TCTGTTG	赤霉素响应元件 Gibberellin-responsive element
AuxRE	TGTCTCAATAAG	生长素响应元件的一部分 Part of an auxin-responsive element
AuxRR-core	GGTCCAT	生长素响应元件 cis-Acting regulatory element involved in auxin responsiveness
ABRE	ACGTG	脱落酸响应元件 cis-Acting element involved in the abscisic acid responsiveness
P-box	CCTTTTG	赤霉素响应元件 Gibberellin-responsive element
TGACG-motif	TGACG	茉莉酸甲酯响应元件 cis-Acting regulatory element involved in the MeJA-responsiveness
TGA-element	AACGAC	生长素响应元件 Auxin-responsive element
TATC-box	TATCCCA	赤霉素响应元件 cis-Acting element involved in gibberellin-responsiveness
circadian	CAAAGATATC	响应昼夜节律控制元件 cis-Acting regulatory element involved in circadian control
AACA_motif	TAACAAACTCCA	胚乳特异性负表达元件 Involved in endosperm-specific negative expression
AT-rich element	ATAGAAATCAA	AT富集DNA结合蛋白位点（ATBP-1） Binding site of AT-rich DNA binding protein（ATBP-1）
CAT-box	GCCACT	分生组织表达相关元件 cis-Acting regulatory element related to meristem expression
GCN4_motif	TGAGTCA	胚乳表达响应元件 cis-Regulatory element involved in endosperm expression
HD-Zip 1	CAAT(A/T)ATTG	栅栏叶肉细胞分化响应元件 Element involved in differentiation of the palisade mesophyll cells
HD-Zip 3	GTAAT(G/C)ATTAC	蛋白结合位点 Protein binding site
MSA-like	TCCAACGGT	细胞周期调节元件cis-Acting element involved in cell cycle regulation
O2-site	GATGATGTGG	玉米醇溶蛋白代谢调节元件cis-Acting regulatory element involved in zein metabolism regulation
RY-element	CATGCATG	种子特异调控元件 cis-Acting regulatory element involved in seed-specific regulation
Unnamed_1	GAATTTAATTAA	60K蛋白结合位点 60K protein binding site
DRE	TACCGACAT	脱水、低温、盐响应元件 cis-Acting element involved in dehydration，low-temp，salt stresses
CCAAT-box	CAACGG	HvMYB1结合位点 HvMYB1 binding site
GC-motif	CCCCCG	增强缺氧特异性诱导元件 Enhancer-like element involved in anoxic specific inducibility
LTR	CCGAAA	低温响应元件 cis-Acting element involved in low-temperature responsiveness
MBS	CAACTG	MYB结合位点参与干旱诱导 MYB binding site involved in drought-inducibility
TCA-element	CCATCTTTTT	水杨酸响应元件 cis-Acting element involved in salicylic acid responsiveness
TC-rich repeats	GTTTTCTTAC	响应防御和应激反应元件 cis-Acting element involved in defense and stress responsiveness
ARE	AAACCA	厌氧诱导关键元件 cis-Acting regulatory element essential for the anaerobic induction
WUN-motif	AAATTTCCT	干旱响应元件Wound-responsive element
CAAT-box	CCAAT	启动子增强子区域元件Common cis-acting element in promoter and enhancer regions
TATA-box	TATA	启动子核心元件Core promoter element around -30 of transcription start

分类	比对结果	功能注释		NCBI登录号
Classify	Alignment result	Functional annotation		GenBank ID
信号转导 Signal transduc- tion	NADH脱氢酶1 α亚复合体亚基1 NADH dehydrogenase 1 alpha subcomplex subunit 1	将电子从NADH转移到线粒体膜呼吸链 Transfer of electrons from NADH to the respiratory chain		XM_029089752.1
	空泡分选受体3类似物 Vacuolar-sorting receptor 3-like	参与网格蛋白包被的囊泡从高尔基体到空泡的分选 Sorting of clathrin-coated vesicles from golgi bodies to vacuoles		XM_008386294.3
	细胞色素b-c1复合体亚基4 Cytochrome b-c1 complex subunit Rieske-4	驱动线粒体氧化磷酸化 Drives mitochondrial oxidative phosphorylation		XR_003767947.1
	细胞色素c氧化酶亚基5C-3 Cytochrome c oxidase subunit 5C-3	细胞色素c氧化酶的核编码多肽链之一，是线粒体电子传递的末端氧化酶 One of the nucleus-coding polypeptide chains of cytochrome C oxidase，the terminal oxidase of mitochondrial electron transport		XM_008370823.3
	钾离子通道AKT2/3 Potassium channel AKT2/3	参与韧皮部汁液中输入和卸载钾的作用 Involved in both loading and unloading potassium into/from the phloem sap		XM_029107389.1
	三酰甘油脂肪酶1 Triacylglycerol lipase 1	催化三酰基甘油的水解 Catalyzes the hydrolysis of triacylglycerol		XM_008340013.3
	硫氧还原蛋白AAED1类似物 Thioredoxin-like protein AAED1	抗氧化活性 Antioxidant activity		XM_008343896.3
	泛酸酯激酶 At2g17340-like		催化泛酸的磷酸化 Catalyzes the phosphorylation of pantothenate	XM_008384280.3
	非典型N-乙酰转移酶p20类似物 Uncharacterized N-acetyltransferase p20-like		参与N-乙酰转移酶活性 N-acetyltransferase activity	XM_029096249.1
转录调控 Transcrip- tional regulation	核糖核酸内切酶YBEY Endoribonuclease YBEY		参与晚期70S核糖体质量控制和16S rRNA 3′端成熟 Involved in late-stage 70S ribosome quality control and in maturation of the 3′ terminus of the 16S rRNA	XM_029106325.1
转录调控 Transcrip- tional regulation	60S核糖体蛋白L10类似物 60S ribosomal protein L10-like		在核糖体的形成中起积极作用 An active role in the formation of ribosomes	XM_029102547.1
抗逆防御 Resistance and defense	脂肪酶PAD4类似物 Lipase-like PAD4		促进植物对入侵性生物营养病原体的先天免疫和识别微生物相关分子模式防御机制 Contributing to the plant innate immunity against invasive biotrophic pathogens and to defense mechanisms upon recognition of microbe-associated molecular patterns	XM_029094810.1
	应激反应蛋白NST1类似物 Stress response protein NST1-like		与MSL1一起作为耐盐性的负调节因子 Negative regulator of salt tolerance along with MSL1	XM_008394867.3
	非特异性脂质转运蛋白3 Non-specific lipid-transfer protein 3		可以跨膜转移磷脂和半乳糖脂，参与植株对生物与非生物胁迫的抗性 Phospholipids and galactose lipids can be transferred across the membrane to participate in plant resistance to biological and abiotic stress	XM_008371792.3
	解毒蛋白35类似物 Protein DETOXIFICATION 35-like		参与类黄酮代谢的多药物和毒素外排转运体 Multidrug and toxin efflux transporters involved in flavonoid metabolism	XM_008381794.3
	单链DNA结合蛋白WHY1 Single stranded DNA binding protein WHY1		致病相关基因PR-10a的转录激活因子 Transcriptional activator of the pathogenesis-related gene PR-10a	NM_001294042.1
	脂质转移蛋白VAS类似物 Lipid transfer-like protein VAS		可以跨膜转移磷脂和半乳糖脂，参与植株对生物与非生物胁迫的抗性 Phospholipids and galactose lipids can be transferred across the membrane to participate in plant resistance to biological and abiotic stress	XM_008342321.3
生长发育 Grow and develop- ment	MADS-box蛋白 SOC1类似物 MADS-box protein SOC1-like		调控开花时间的转录因子 Transcription factors that regulate flowering time	XM_008378604.3
生长发育 Grow and develop- ment	质体转录活性蛋白7 Plastid transcriptionally active 7		参与质体基因表达，是类囊体形成所必需的 Involved in plastid gene expression，essential for thylakoid formation	XM_008382937.3
	核酸外切酶DPD1 Exonuclease DPD1		花粉发育过程中细胞器DNA降解所需的外切酶 Exonuclease enzymes required for degradation of organelle DNA during pollen development	XM_017336537.2
	光合作用NDH亚基的腔定位5 Photosynthetic NDH subunit of lumenal location 5		叶绿体NAD(P)H脱氢酶复合物形成和活性所必需 Essential for chloroplast NAD(P)H dehydrogenase complex formation and activity	XM_029091434.1
	锌指蛋白CONSTANS-LIKE13类似物 Zinc finger protein CONSTANS-LIKE 13-like		促进开花 Accelerate bloom	XM_029100386.1
	二氨基庚二酸表异构酶 Diaminopimelate epimerase		参与L-赖氨酸生物合成和肽聚糖中五肽的生物合成 Involved in the L-lysine biosynthesis and in the biosynthesis of the pentapeptide incorporated in the peptidoglycan moiety	XM_029101158.1
	内质网素同源物 Endoplasmin homolog		与芽、根和花分生组织形成及花粉管伸长有关 Related to shoot apical meristem，root apical meristem floral meristem formation and pollen tube elongation	XM_009336652.2
	碳酸酐酶2类似物 Carbonic anhydrase 2-like		催化二氧化碳的可逆水合作用形成碳酸氢盐 Catalyzes the reversible hydration of carbon dioxide to form bicarbonate	XM_008359520.3
	叶绿体 Chloroplast		从细胞质向叶绿体输入核编码的光合前蛋白 Nucleo-coded prephotosynthetic proteins are imported from the cytoplasm to chloroplasts	OK545754.1
	LRR受体样丝氨酸/苏氨酸蛋白激酶 LRR receptor-like serine/threonine-protein kinase		花粉发育的关键调控因子 Key regulator of anther development	XM_029088102.1
	生育酚-O-甲基转移酶 Tocopherol O-methyltransferase		参与生育酚的合成 Involved in the synthesis of tocopherol	XM_029092547.1
	脂肪酸去饱和酶4 Fatty acid desaturase 4		参与生产含有叶绿体特异性的磷脂酰甘油 Involved in the production of chloroplast-specific phosphatidylglycerol	XM_008373042.2

分类	比对结果	功能注释		NCBI登录号
Classify	Alignment result	Functional annotation		GenBank ID
信号转导 Signal transduc- tion	NADH脱氢酶1 α亚复合体亚基1 NADH dehydrogenase 1 alpha subcomplex subunit 1	将电子从NADH转移到线粒体膜呼吸链 Transfer of electrons from NADH to the respiratory chain		XM_029089752.1
	空泡分选受体3类似物 Vacuolar-sorting receptor 3-like	参与网格蛋白包被的囊泡从高尔基体到空泡的分选 Sorting of clathrin-coated vesicles from golgi bodies to vacuoles		XM_008386294.3
	细胞色素b-c1复合体亚基4 Cytochrome b-c1 complex subunit Rieske-4	驱动线粒体氧化磷酸化 Drives mitochondrial oxidative phosphorylation		XR_003767947.1
	细胞色素c氧化酶亚基5C-3 Cytochrome c oxidase subunit 5C-3	细胞色素c氧化酶的核编码多肽链之一，是线粒体电子传递的末端氧化酶 One of the nucleus-coding polypeptide chains of cytochrome C oxidase，the terminal oxidase of mitochondrial electron transport		XM_008370823.3
	钾离子通道AKT2/3 Potassium channel AKT2/3	参与韧皮部汁液中输入和卸载钾的作用 Involved in both loading and unloading potassium into/from the phloem sap		XM_029107389.1
	三酰甘油脂肪酶1 Triacylglycerol lipase 1	催化三酰基甘油的水解 Catalyzes the hydrolysis of triacylglycerol		XM_008340013.3
	硫氧还原蛋白AAED1类似物 Thioredoxin-like protein AAED1	抗氧化活性 Antioxidant activity		XM_008343896.3
	泛酸酯激酶 At2g17340-like		催化泛酸的磷酸化 Catalyzes the phosphorylation of pantothenate	XM_008384280.3
	非典型N-乙酰转移酶p20类似物 Uncharacterized N-acetyltransferase p20-like		参与N-乙酰转移酶活性 N-acetyltransferase activity	XM_029096249.1
转录调控 Transcrip- tional regulation	核糖核酸内切酶YBEY Endoribonuclease YBEY		参与晚期70S核糖体质量控制和16S rRNA 3′端成熟 Involved in late-stage 70S ribosome quality control and in maturation of the 3′ terminus of the 16S rRNA	XM_029106325.1
转录调控 Transcrip- tional regulation	60S核糖体蛋白L10类似物 60S ribosomal protein L10-like		在核糖体的形成中起积极作用 An active role in the formation of ribosomes	XM_029102547.1
抗逆防御 Resistance and defense	脂肪酶PAD4类似物 Lipase-like PAD4		促进植物对入侵性生物营养病原体的先天免疫和识别微生物相关分子模式防御机制 Contributing to the plant innate immunity against invasive biotrophic pathogens and to defense mechanisms upon recognition of microbe-associated molecular patterns	XM_029094810.1
	应激反应蛋白NST1类似物 Stress response protein NST1-like		与MSL1一起作为耐盐性的负调节因子 Negative regulator of salt tolerance along with MSL1	XM_008394867.3
	非特异性脂质转运蛋白3 Non-specific lipid-transfer protein 3		可以跨膜转移磷脂和半乳糖脂，参与植株对生物与非生物胁迫的抗性 Phospholipids and galactose lipids can be transferred across the membrane to participate in plant resistance to biological and abiotic stress	XM_008371792.3
	解毒蛋白35类似物 Protein DETOXIFICATION 35-like		参与类黄酮代谢的多药物和毒素外排转运体 Multidrug and toxin efflux transporters involved in flavonoid metabolism	XM_008381794.3
	单链DNA结合蛋白WHY1 Single stranded DNA binding protein WHY1		致病相关基因PR-10a的转录激活因子 Transcriptional activator of the pathogenesis-related gene PR-10a	NM_001294042.1
	脂质转移蛋白VAS类似物 Lipid transfer-like protein VAS		可以跨膜转移磷脂和半乳糖脂，参与植株对生物与非生物胁迫的抗性 Phospholipids and galactose lipids can be transferred across the membrane to participate in plant resistance to biological and abiotic stress	XM_008342321.3
生长发育 Grow and develop- ment	MADS-box蛋白 SOC1类似物 MADS-box protein SOC1-like		调控开花时间的转录因子 Transcription factors that regulate flowering time	XM_008378604.3
生长发育 Grow and develop- ment	质体转录活性蛋白7 Plastid transcriptionally active 7		参与质体基因表达，是类囊体形成所必需的 Involved in plastid gene expression，essential for thylakoid formation	XM_008382937.3
	核酸外切酶DPD1 Exonuclease DPD1		花粉发育过程中细胞器DNA降解所需的外切酶 Exonuclease enzymes required for degradation of organelle DNA during pollen development	XM_017336537.2
	光合作用NDH亚基的腔定位5 Photosynthetic NDH subunit of lumenal location 5		叶绿体NAD(P)H脱氢酶复合物形成和活性所必需 Essential for chloroplast NAD(P)H dehydrogenase complex formation and activity	XM_029091434.1
	锌指蛋白CONSTANS-LIKE13类似物 Zinc finger protein CONSTANS-LIKE 13-like		促进开花 Accelerate bloom	XM_029100386.1
	二氨基庚二酸表异构酶 Diaminopimelate epimerase		参与L-赖氨酸生物合成和肽聚糖中五肽的生物合成 Involved in the L-lysine biosynthesis and in the biosynthesis of the pentapeptide incorporated in the peptidoglycan moiety	XM_029101158.1
	内质网素同源物 Endoplasmin homolog		与芽、根和花分生组织形成及花粉管伸长有关 Related to shoot apical meristem，root apical meristem floral meristem formation and pollen tube elongation	XM_009336652.2
	碳酸酐酶2类似物 Carbonic anhydrase 2-like		催化二氧化碳的可逆水合作用形成碳酸氢盐 Catalyzes the reversible hydration of carbon dioxide to form bicarbonate	XM_008359520.3
	叶绿体 Chloroplast		从细胞质向叶绿体输入核编码的光合前蛋白 Nucleo-coded prephotosynthetic proteins are imported from the cytoplasm to chloroplasts	OK545754.1
	LRR受体样丝氨酸/苏氨酸蛋白激酶 LRR receptor-like serine/threonine-protein kinase		花粉发育的关键调控因子 Key regulator of anther development	XM_029088102.1
	生育酚-O-甲基转移酶 Tocopherol O-methyltransferase		参与生育酚的合成 Involved in the synthesis of tocopherol	XM_029092547.1
	脂肪酸去饱和酶4 Fatty acid desaturase 4		参与生产含有叶绿体特异性的磷脂酰甘油 Involved in the production of chloroplast-specific phosphatidylglycerol	XM_008373042.2