Analysis of different expressiongenes between cross-compatibility and cross-incompatibility within pistils of Lilium spp
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摘要: 东方百合与卷瓣组野生百合杂交(简称OS组合)，属于亲缘关系最远的组间杂交，杂交障碍最大。挖掘雌蕊中杂交亲和性与不亲和性差异表达基因，有可能解释不亲和性杂交中引起花粉管定向生长异常的机理，进而阐明不亲和性杂交的分子机理，为实施克服杂交不亲和障碍技术措施提供依据。本研究采用抑制消减杂交技术(SSH技术)，分别以不亲和性杂交和亲和性杂交的东方百合雌蕊为试验组(tester)和驱动组(driver)，建立正向抑制消减杂交文库，随机挑选180个阳性克隆，经过测序、去劣、去冗余、序列比对，有113个EST与数据库中的已知序列具有同源性，最终获得10个差异表达基因。采用RT-PCR技术对10个差异表达基因进行验证，其中有1个基因在不亲和性杂交的雌蕊中表达上调、7个基因表达下调、2个基因无明显变化。对差异表达基因进行功能注释及分类，差异表达基因主要集中于信号转导(包括丝氨酸/苏氨酸蛋白磷酸酶PP2A-3、钙依赖蛋白激酶CDPK、小G结合蛋白)、抗逆防御(包括分子伴侣(clpB)、过氧化氢酶CAT2)、转运(包括质膜ATPase、焦磷酸酶质子泵)、蛋白命运(包括60S核糖体蛋白)等功能类别。据此推测，在东方百合×山丹组(系)间不亲合性杂交育种过程中，可能引起雌蕊与花粉管信号交流异常、物质转运能力减弱和雌蕊耐受逆境能力降低，从而导致杂交不亲和性。Abstract: The hybridization of Lilium Oriental hybrids and wild lily of Sect. Sinomartagon Comber is the farmost genetic relationship and it's difficult to achieve. Researching the different expression genes in pistil of cross-compatibility and cross-incompatibility may explain the mechanism of causing orientated growth of pollen tube and the molecular mechanism of incompatible hybridization, and provide the basis for overcoming cross-incompatibility. We constructed a forward suppression subtractive hybridization (SSH) library by using the cross-incompatible Lilium Oriental hybrid pistils as the tester, and the cross-compatible ones as the driver. 180 positive clones were randomly screened out. Through sequencing, deleting the bad, removing the redundant and aligning the sequences, 113 ESTs were found to be homologous with known sequences in the bioinformatics databases. RT-PCR was performed to test the 10 selected different expression genes, which showed 1 gene was up-regulated, 7 genes were down-regulated and 2 genes were almost unchanged. Gene annotation and functional category analysis showed that the different expression genes mainly contained signal transduction(including PP2A-3, CDPK and small GTP-binding protein), transport-related(including clpB and CAT2), stress/defense(including ATPase and pyrophosphatase proton pump), protein fate related genes(including 60S ribosomal protein). It was speculated that the incompatibility in the section cross of Oriental hybrids ×Lilium pumilum was probably resulted from the abnormal signal communication between pistils and pollens, the deduced substance transport ability in pistils, the weaker stress resistance ability in pistils after incompatible hybridization.
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图 1 总RNA和mRNA电泳检测
M. DL 200 marker; T.试验组; D.驱动组。
Figure 1. Electrophoresis detection of total RNA and mRNA
M, DL 200 marker; T, Tester; D, Driver.
图 2 cDNA酶切前后的电泳图
M. DL 2 000 marker; T1.试验组cDNA; T2.酶切后试验组cDNA; D1.驱动组cDNA; D2.酶切后试验组cDNA。
Figure 2. Electrophoresis pattern of cDNA before and after digestion
M, DL 2 000 marker; T1, Tester cDNA; T2, Tester cDNA after digestion; D1, Driver cDNA; D2, Driver cDNA after digestion.
图 3 二次PCR产物电泳图
M.DL 2 000 marker；E1.正向消减第一次PCR产物; 1-c.正向未消减第一次PCR产物；pE1.正向消减第二次PCR产物; p1-c.正向未消减第二次PCR产物。
Figure 3. Electrophoresis result of the first and second PCR
M, DL 2 000 marker; E1, first PCR production of positive subtraction; 1-c, first PCR production of positive non-subtraction; pE1, second PCR production of positive subtraction; p1-c, second PCR production of positive non-subtraction.
图 4 消减cDNA文库消减效率分析
M.DL 2 000 marker；1、2、3、4、5分别为消减后第18、23、28、33、36个循环的PCR产物；6、7、8、9、10分别为未消减的第18、23、28、33、36个循环的PCR产物。
Figure 4. Cutting efficiency analysis of subtracted cDNA library
M, DL 2 000 marker; 1, 2, 3, 4, 5 are the PCR production of the 18th, 23rd, 28th, 33rd, 36th cycles after subtracted respectively; 6, 7, 8, 9, 10 are the non-subtractive PCR production of the 18th, 23rd, 28th, 33rd, 36th cycles, respectively.
图 6 试验组和驱动组中差异表达基因的GO分类
细胞cell；细胞组分Cell component；高分子配合物Macromolecular complex；细胞器Organelle；细胞器组分Organelle part；抗氧化剂Antioxidant；结合Binding；催化剂Catalytic；电子载体Electron carrier；结构分子Structural molecule；转录调节因子Transcription regulator；转运Transporter；解剖结构组成Anatomical structure formation；生物调节Biological regulation；细胞组成生物源Cellular component biogenesis；细胞组织组件Cellular component organization；细胞过程Cellular process；建立定位Establishment of localization；定位Localization；代谢过程Metabolic process；色素形成Pigmentation；刺激反应Response to stimulus
Figure 6. GO classification of different expression genes of tester and driver
图 7 基因表达量检测
T.试验组Tester; D.驱动组Driver; 1.柱头Stigma；2.花柱Style；3.子房Ovary；4.花瓣Petal；5.叶Leaf；6.茎Stem；7.鳞茎Bulb
Figure 7. Detection of gene expression level
表 1 特异性引物
Table 1. Specific primer
基因名称
Gene name上游引物(5′-3′)
Forward primer(5′-3′)下游引物(5′-3′)
Reverse primer(5′-3′)退火温度
Annealing
temperature/℃60S核糖体蛋白
60S ribosomal protein(RP)AATCCCTTGAATCCTCTTGCC AGAGAAGGCGAAGATGGTG 54 液泡膜H+-ATPase
Vacuolar H+-ATPase(V-H+-ATPase)AGAGAGAAGATGACCTGAATGAAAT CCAAACGATGCTTGATGACG 54 小G结合蛋白
Small GTP-binding protein (small GTP)ACCTGCTCAAGAACTAGAAG AGTAGGGAACAAAACAAACAC 51 丝氨酸/苏氨酸蛋白磷酸酶
Serine/Threonine-protein phosphatase 2A catalytic subunit3(PP2A-3)GGAGGAAAAGATGAGCGGG AATAATAGCCACGGTCCACATAATC 55 分子伴侣CLPB
Molecular chaperone CLPB(CLPB)TGACTATGCTGTTGATCTGC ATGCAGTTTCGAGATTGAT 49 过氧化氢酶2
Catalase2(CAT2)CAACCTGGAGAGCGATACC GTCGAAGGTAGTAAGCC 50 质膜ATPase
Plasma membrane atpase (PM-ATPase)AGTTTAATGCAAGCGATAT AGCAAGAAGAATTATGGG 48 焦磷酸盐能膜质子泵3
Pyrophosphate-energized membrane
proton pump 3AATCCCTTATTCCACAAACAAG ATCTTCGTTGACCTGGCTAAG 52 肌动蛋白解聚合因子
Actin depolymerizing factor(ADF)GTACCCAAACAAGAAGCACAT TGATGTGTCGAGGGTGAGGAGT 53 钙依赖蛋白激酶Calcium-dependent protein kinase(CDPK) CAGTATTAGAACTCATTGGCAC GTAATCCCCATATTCACTGCTG 54 
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表 2 试验组与驱动组部分差异表达基因分类
Table 2. Classification of partial different expression of tester and driver
功能类别
Functional category基因注释
Gene annotation基因ID
Gene IDE值
E value信号转导相关基因Signal transduction genes 类钙周期素结合蛋白Ealeyelin- binding protein(CacyBP) OS37、OS41 2×10-66,3×10-63 丝氨酸/苏氨酸蛋白磷酸酶Serine/threonine-protein phosphatase 2A catalytic subunit3(PP2A-3) OS52、OSP27 0，2×10-11 ADP核糖基化因子ADP-ribosylation factor(ARF) OS85 3×10-118 富含亮氨酸重复序列类似受体激酶Leucine-rich repeat receptor kinase (LRR-RLKs) OSP4 3×10-73 周期素依赖性蛋白激酶调控基因Cyclindependent kinase(CDK) OSP12 4×10-20 丝氨酸/苏氨酸蛋白磷酸酶2A调节亚基BSerine/threonine-protein phosphatase 2A catalytic subunitB(PP2A-B) OSP19 7×10-baby5 S期激酶相关蛋白1S-phase kinase associated protein(SKP1) OSP32 8×10-55 小G结合蛋白Small GTP-binding protein(small GTP) OSP62 9×10-3 SCA基因Stigma/style Cysteine-rich Adhesin(SCA) OS36 3×10-12 钙依赖蛋白激酶Calcium-dependent protein kinase(CDPK) OSP63 3×10-3 胚胎发育晚期丰富蛋白1Late embryogenesis abundant proteins 1(LEC1) OSP47 3×10-8 转运类相关基因
Transport-related genes焦磷酸盐能膜质子泵3Pyrophosphate-energized membrane proton pump 3 OS33 8×10-73 质膜内在蛋白1Plasma membrane intrinsic proteins 1(PIP1) OS35、OSP21 2×10-62,1×10-9 酰基载体蛋白Acyl carrier protein(ACP) OS55、OS87 4×10-26,4×10-26 ADP/ATP载体蛋白ADP/ATPcarrier protein(AAC) OS59 5×10-25 水通道蛋白2Aquaporin 2(AQP2) OS109 6×10-74 焦磷酸酶/磷酸二酯酶Nucleotide pyrophosphatase/phosphodiesterase1(ENPP) OSPP57 2×10-45 SCA基因Stigma/style Cysteine-rich Adhesin(SCA) OS36 3×10-12 质膜ATPase 4-likePlasma membrane atpase(PM-ATPase 4L) OSPP06 2×10-66 抗逆防御相关基因Stress/defense genes 亲环蛋白基因Cyclophilin(CyP) OS17 5×10-23 半胱氨酸合酶Cysteine synthase(Csase) OS25 4×10-77 过氧化氢酶2Catalase2(CAT2) OS29 1×10-36 分子伴侣CLPB Molecular chaperone CLPB(CLPB) OS65 1×10-107 抗坏血酸过氧化物酶Ascorbate peroxidase(APX) OS69 8×10-89 脂氧合酶lipoxygenase(LOX) OS71 2×10-10 丝氨酸乙醛酸氨基转移酶Rine:Glyoxylateaminotrans-ferase(SGAT) OS74 4×10-175 细胞色素P450类TBP蛋白Cytochrome P450 like TBP(CYP-TBP) OS82 0 ACC氧化酶ACC oxidase(ACO) OS88 4×10-180 脱水蛋白Dehydrin(DHN) OS90 2×10-5 液泡加工酶3Vacuolar processing enzyme(VPE3) OS95 3×10-60 硫氧还原蛋白Thioredoxin(TRX) OS108 5×10-50 谷胱甘肽过氧化物酶Glutathione peroxidase(GSH-Px) OSP59 2×10-19 金属硫蛋白Metallothionein(MT) OS3、OSP38 2×10-113、3×10-64 蛋白质命运相关基因Protein fate related genes 转录起始因子IIB-like Transcription initiation factor IIB-like(IIB-L) OS119 4×10-42 起始因子iso-4F Eukaryotic initiation factor 4F (eIF-(iso)4F) OS67 4×10-17 翻译起始因子eIF-5A前体蛋白Translation initiation factor 5A precursor(ec-eIF5A) OSP50 6×10-66 核糖体蛋白L2 Ribosomal protein(rp-L2) OS6、OS96 2×10-10,3×10-6 60S核糖体蛋白60S ribosomal protein(RP) OSP7、OSP16 5×10-87,5×10-51
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