番茄花柄脱落相关基因LelDL和LeHAESA克隆功能验证及LeMKKs和LeMPKs的钙素调控
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摘要
器官脱落（abscission）是植物细胞、组织或器官脱离母体的生理过程，器官脱落发生的特定的区域叫做离区（abscission zone）。拟南芥中，大多数影响花器官脱落的基因会不同程度的提前或延迟脱落，目前已经明确的花器官脱落的信号转导途径是BOP1/BOP2→AtIDA→AtHAESA→AtMKKK?→AtMKK4/MKK5→AtMPK3/MPK6→花器官脱落。虽然目前已经分离得到一些与离区发育相关的基因，但对番茄花柄脱落的信号转导途径的研究较少。为了阐明番茄花柄脱落的信号转导机制，本研究主要结果如下：
1分别利用LeIDL和LeHAESA中的保守区域设计简并引物。通过同源扩增的方法分别获得400bp和700bp左右的的与其他植物IDA和HAESA相似的核心序列，再用RACE(cDNA末端快速扩增技术)得到3’端和5’端序列，进行序列拼接，最终获得两条长度为510bp和3035bp的序列。分别将其命名为LeIDL和LeHAESA.生物信息学表明：它们分别具有510bp和2667bp的ORF框，与已报道的多个的IDA、IDL（IDA-LIKE）和HAESA、HSL（HAESA-LIKE）具有较高的相似度。推断其编码101个和888个氨基酸，分子量分别为11456.3和97874.1,等电点分别为9.56和6.35。qRT-PCR分析在花的不同发育阶段，这两个基因在花柄离区具有表达，不同的是LeIDL在整个花发育过程中表达较为稳定。随着花发育进程的进行，LeHAESA呈现出明显的规律性:在花瓣未打开前，其表达量与花发育呈正相关。在开花后（果实不同发育阶段），LeHAESA在坐果初期表达量较高，随后下降，直至过程晚熟，表达量再次上升。在花的不同组织中，LeIDL在花瓣表达很高其次是雄蕊，LeHAESA在雌蕊的表达量最高，其次是雄蕊。这说明不仅参与了番茄花柄的脱落同时也参与的番茄花的发育。
2本试验以“中蔬六号”番茄作为试材，研究了生长素IAA与细胞分裂素6-BA对其愈伤组织的产生和不定芽的诱导，细胞分裂素NAA对不定根的诱导。经研究认为，“中蔬六号”最佳诱导分化培养基为MS+0.2mg·L-1IAA+1.0mg·L-16-BA,最佳诱导生根培养基配方为MS+0.05mg·L-1NAA。同时以p MDC141为过表达载体，研究了转化细胞对Hyp抗性进行了0mgL-1、5mgL-1、7mgL-1、9mgL-1、11mgL-1、13mgL-1等6个梯度试验，浓度达到9mgL-1外植体伤口处褐化，直至完全死亡。因此后续试验不定芽抗性筛选浓度均采用7mgL-1。并获得8株阴性对照。以PB7GWIWG(II)为RNAi表达载体，研究了转化细胞对PPT抗性，初次筛选选用0mgL-1、0.5mgL-1、1.0mgL-1、1.5mgL-1、2.0mgL-12.5mgL-1等6个梯度，再次筛选在浓度为0mgL-1，0.6mgL-1、0.8mgL-1、1.0mgL-1、1.2mgL-1、1.4mgL-1PPT，当PPT浓度为1.0mgL-1时，极少有芽分化，并且分化出的芽逐渐变黄至白化。本试验以后的不定芽抗性筛选浓度均采用0.8mgL-1
3利用Gateway技术构建分别构建LeIDL和LeHAESA基因过表达和RNAi表达载体，并成功转化到番茄中，得到的转基因植株进行PCR验证，结果发现：转基因植株中过表达技术提高了转基因植株中LeIDL和LeHAESA的转录水平，RNAi技术降低了LeIDL和LeHAESA基因的表达。35S: LeIDL植株的花序数目增多；RNAi:LeIDL转基因果实发育异常、心室异常增多；35S:LeHAESA株系中花器官、果实异常，同时叶色发生改变。
4系统系统发生树表明LeMKK2与AtMKK4/AtMKK5， LeMPK1、 LeMPK2与AtMPK6，LeMPK3与AtMPK6均具有较近的亲缘关系，进而推测它们具有相似的的功能。qRT-PCR结果说明LeMKK2、LeMPK1、LeMPK2正调控花柄脱落，钙在脱落初期就完成对MAP级联途径的信号的放大作用。钙和W7共同处理Ca+W7恢复或部分恢复了Ca对LeMKK1,3,4基因表达的抑制作用，但却加剧了对LeMKK2的抑制作用，完全恢复了Ca对LeMPK1的抑制作用。
Abscission is a physiological process that plant cell, tissue or organ separate from thematernal. The specific region abscission occurred at is called abscission zone. In tomatoproduction, especially in the facilities tomato cultivation in winter and spring, it easily causesflower and fruit dropping. Although some genes related to the abscission development havebeen isolated, less study about the signal transduction pathways of tomato pedicel abscissionhave been done. In this article, tomato pedicels were taken as test materials, cDNA sequencewas gained by RACE and the application of reverse genetics made detailed analysis on them.The main results were as follows:
1. The primers were designed according to IDA and HAESA conserved regions. About400bp and700bp core sequence similar with other plants were obtained by homologousamplification. Then we gained the3'and5' terminal by using RACE(cDNA ends rapidamplification technology). Finally510bp and3035bp were obtained by splice, named themLeIDL and LeHAESA Respectively. Bioinformatics analysis explain that: They have510bpand2667bp ORF respectively, appear high similarity with IDA, IDL（IDA-LIKE）, HAESA,HSL（HAESA-LIKE） as reported. Infering they encording101and888amino acids, withmolecular weights of11456.3and97874.1, isoelectric point of9.56and6.35repectively. Thetwo genes have expressed at pedicel abscission zone in different developmental stages of theflower by qRT-PCR analysis, but LeIDL was more stablely expressed during the wholeprocess. With the process of flower development, LeHAESA showed obvious regularity:while the petals had not open, its expression was positively correlated with its development.After flowering, it showed high expression level at early fruit set and then decreased until latematuring, it rose again. In different tissues of the flower, LeIDL expressed highest in petalsand stamens second. While LeHAESA expressed highest in pistil and followed by stamens.The results shows that they involved in both tomato pedicel abscission and flowersdevelopment.
2. In this study,“Chinese vegetables6th” was taken as the test material to Investigate theinduced role of IAA and6-BA in callus and adventitious buds and NAA in adventitious roots.The results showed the best induced differentiation medium formula for “Chinese vegetables6th” is MS+0.2mg·L-1IAA+1.0mg·L-16-BA, and the best induced rooting medium formulais MS+0.05mg·L-1NAA.In this study, pMDC141was taken as overexpression vector. Inorder to find the transformed cells resistance against Hyp, we set six-gradient test as0mgL-1,5mgL-1,7mgL-1,9mgL-1,11mgL-1,13mgL-1. Once it reached9mgL-1, explant woundappeared brown until it completely dead. As a result,7mgL-1was used in the follow-up testsof adventitious buds resistance screening concentration, and eight negative control were
obtained. At the same time, PB7GWIWG(II) was taken as RNAi expression vector toinvestigate the transformed cells resistance against PPT. In initial screening, we setsix-gradient as0mgL-1,0.5mgL-1,1.0mgL-1,1.5mgL-1,2.0mgL-1,2.5mgL-1and inlater screening, we set six-gradient as0mgL-1,0.6mgL-1,0.8mgL-1,1.0mgL-1,1.2mgL-11.4mgL-1PPT. When the concentration of PPT was1.0mgL-1, there was few budsdifferentiation, and moreover the buds turned yellow to bleaching gradually. Therefore,0.8mgL-1was used in the follow-up tests of adventitious buds resistance screeningconcentration.
3Buliding expression vector of LeIDL and LeHAESA using Gateway technology. Thetransgenic plants were tested by PCR.The results found that the transcription was improvedby technology of overexpression, the transcription; was inhibited by RNAi. Plantinflorescence are added in35S: LeIDL; the development of fruit was anomaly in RNAi:LeIDL; floral organs anf fruit was unnormal in35S:LeHAESA
4System phylogenetic tree showed that LeMKK2and AtMKK4/AtMKK5,LeMPK1,LeMPK2and AtMPK6, LeMPK3and AtMPK6are close relatives, suggesting that they havesimilar function. qRT-PCR analysis showed LeMKK2, LeMPK1, LeMPK2positively regulatepedicel abscission. Ca play an amplification role of the signal cascades of MAP in the earlytime of abscission. The treatment of Ca+W7restored or partly restored the inhibitory effect ofCa on LeMKK1,3,4gene expression, but exacerbated the inhibitory effect on LeMKK2andthe treatment completely restored the inhibition effect of Ca on LeMPK1.
1分别利用LeIDL和LeHAESA中的保守区域设计简并引物。通过同源扩增的方法分别获得400bp和700bp左右的的与其他植物IDA和HAESA相似的核心序列，再用RACE(cDNA末端快速扩增技术)得到3’端和5’端序列，进行序列拼接，最终获得两条长度为510bp和3035bp的序列。分别将其命名为LeIDL和LeHAESA.生物信息学表明：它们分别具有510bp和2667bp的ORF框，与已报道的多个的IDA、IDL（IDA-LIKE）和HAESA、HSL（HAESA-LIKE）具有较高的相似度。推断其编码101个和888个氨基酸，分子量分别为11456.3和97874.1,等电点分别为9.56和6.35。qRT-PCR分析在花的不同发育阶段，这两个基因在花柄离区具有表达，不同的是LeIDL在整个花发育过程中表达较为稳定。随着花发育进程的进行，LeHAESA呈现出明显的规律性:在花瓣未打开前，其表达量与花发育呈正相关。在开花后（果实不同发育阶段），LeHAESA在坐果初期表达量较高，随后下降，直至过程晚熟，表达量再次上升。在花的不同组织中，LeIDL在花瓣表达很高其次是雄蕊，LeHAESA在雌蕊的表达量最高，其次是雄蕊。这说明不仅参与了番茄花柄的脱落同时也参与的番茄花的发育。
2本试验以“中蔬六号”番茄作为试材，研究了生长素IAA与细胞分裂素6-BA对其愈伤组织的产生和不定芽的诱导，细胞分裂素NAA对不定根的诱导。经研究认为，“中蔬六号”最佳诱导分化培养基为MS+0.2mg·L-1IAA+1.0mg·L-16-BA,最佳诱导生根培养基配方为MS+0.05mg·L-1NAA。同时以p MDC141为过表达载体，研究了转化细胞对Hyp抗性进行了0mgL-1、5mgL-1、7mgL-1、9mgL-1、11mgL-1、13mgL-1等6个梯度试验，浓度达到9mgL-1外植体伤口处褐化，直至完全死亡。因此后续试验不定芽抗性筛选浓度均采用7mgL-1。并获得8株阴性对照。以PB7GWIWG(II)为RNAi表达载体，研究了转化细胞对PPT抗性，初次筛选选用0mgL-1、0.5mgL-1、1.0mgL-1、1.5mgL-1、2.0mgL-12.5mgL-1等6个梯度，再次筛选在浓度为0mgL-1，0.6mgL-1、0.8mgL-1、1.0mgL-1、1.2mgL-1、1.4mgL-1PPT，当PPT浓度为1.0mgL-1时，极少有芽分化，并且分化出的芽逐渐变黄至白化。本试验以后的不定芽抗性筛选浓度均采用0.8mgL-1
3利用Gateway技术构建分别构建LeIDL和LeHAESA基因过表达和RNAi表达载体，并成功转化到番茄中，得到的转基因植株进行PCR验证，结果发现：转基因植株中过表达技术提高了转基因植株中LeIDL和LeHAESA的转录水平，RNAi技术降低了LeIDL和LeHAESA基因的表达。35S: LeIDL植株的花序数目增多；RNAi:LeIDL转基因果实发育异常、心室异常增多；35S:LeHAESA株系中花器官、果实异常，同时叶色发生改变。
4系统系统发生树表明LeMKK2与AtMKK4/AtMKK5， LeMPK1、 LeMPK2与AtMPK6，LeMPK3与AtMPK6均具有较近的亲缘关系，进而推测它们具有相似的的功能。qRT-PCR结果说明LeMKK2、LeMPK1、LeMPK2正调控花柄脱落，钙在脱落初期就完成对MAP级联途径的信号的放大作用。钙和W7共同处理Ca+W7恢复或部分恢复了Ca对LeMKK1,3,4基因表达的抑制作用，但却加剧了对LeMKK2的抑制作用，完全恢复了Ca对LeMPK1的抑制作用。
Abscission is a physiological process that plant cell, tissue or organ separate from thematernal. The specific region abscission occurred at is called abscission zone. In tomatoproduction, especially in the facilities tomato cultivation in winter and spring, it easily causesflower and fruit dropping. Although some genes related to the abscission development havebeen isolated, less study about the signal transduction pathways of tomato pedicel abscissionhave been done. In this article, tomato pedicels were taken as test materials, cDNA sequencewas gained by RACE and the application of reverse genetics made detailed analysis on them.The main results were as follows:
1. The primers were designed according to IDA and HAESA conserved regions. About400bp and700bp core sequence similar with other plants were obtained by homologousamplification. Then we gained the3'and5' terminal by using RACE(cDNA ends rapidamplification technology). Finally510bp and3035bp were obtained by splice, named themLeIDL and LeHAESA Respectively. Bioinformatics analysis explain that: They have510bpand2667bp ORF respectively, appear high similarity with IDA, IDL（IDA-LIKE）, HAESA,HSL（HAESA-LIKE） as reported. Infering they encording101and888amino acids, withmolecular weights of11456.3and97874.1, isoelectric point of9.56and6.35repectively. Thetwo genes have expressed at pedicel abscission zone in different developmental stages of theflower by qRT-PCR analysis, but LeIDL was more stablely expressed during the wholeprocess. With the process of flower development, LeHAESA showed obvious regularity:while the petals had not open, its expression was positively correlated with its development.After flowering, it showed high expression level at early fruit set and then decreased until latematuring, it rose again. In different tissues of the flower, LeIDL expressed highest in petalsand stamens second. While LeHAESA expressed highest in pistil and followed by stamens.The results shows that they involved in both tomato pedicel abscission and flowersdevelopment.
2. In this study,“Chinese vegetables6th” was taken as the test material to Investigate theinduced role of IAA and6-BA in callus and adventitious buds and NAA in adventitious roots.The results showed the best induced differentiation medium formula for “Chinese vegetables6th” is MS+0.2mg·L-1IAA+1.0mg·L-16-BA, and the best induced rooting medium formulais MS+0.05mg·L-1NAA.In this study, pMDC141was taken as overexpression vector. Inorder to find the transformed cells resistance against Hyp, we set six-gradient test as0mgL-1,5mgL-1,7mgL-1,9mgL-1,11mgL-1,13mgL-1. Once it reached9mgL-1, explant woundappeared brown until it completely dead. As a result,7mgL-1was used in the follow-up testsof adventitious buds resistance screening concentration, and eight negative control were
obtained. At the same time, PB7GWIWG(II) was taken as RNAi expression vector toinvestigate the transformed cells resistance against PPT. In initial screening, we setsix-gradient as0mgL-1,0.5mgL-1,1.0mgL-1,1.5mgL-1,2.0mgL-1,2.5mgL-1and inlater screening, we set six-gradient as0mgL-1,0.6mgL-1,0.8mgL-1,1.0mgL-1,1.2mgL-11.4mgL-1PPT. When the concentration of PPT was1.0mgL-1, there was few budsdifferentiation, and moreover the buds turned yellow to bleaching gradually. Therefore,0.8mgL-1was used in the follow-up tests of adventitious buds resistance screeningconcentration.
3Buliding expression vector of LeIDL and LeHAESA using Gateway technology. Thetransgenic plants were tested by PCR.The results found that the transcription was improvedby technology of overexpression, the transcription; was inhibited by RNAi. Plantinflorescence are added in35S: LeIDL; the development of fruit was anomaly in RNAi:LeIDL; floral organs anf fruit was unnormal in35S:LeHAESA
4System phylogenetic tree showed that LeMKK2and AtMKK4/AtMKK5,LeMPK1,LeMPK2and AtMPK6, LeMPK3and AtMPK6are close relatives, suggesting that they havesimilar function. qRT-PCR analysis showed LeMKK2, LeMPK1, LeMPK2positively regulatepedicel abscission. Ca play an amplification role of the signal cascades of MAP in the earlytime of abscission. The treatment of Ca+W7restored or partly restored the inhibitory effect ofCa on LeMKK1,3,4gene expression, but exacerbated the inhibitory effect on LeMKK2andthe treatment completely restored the inhibition effect of Ca on LeMPK1.
引文
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