内源茉莉酸（JA）在水稻颖花开放中的作用
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摘要
水稻颖花开放是一高度协调的发育过程,包括内外稃的张开、花丝伸长和花药开裂,受到遗传因素和环境因素(天气和人工处理等)的影响。颖花开放调控机制的研究有助于解决水稻开花期高温引起的颖花不育以及杂交水稻制种中经常出现的父母本花时不遇的问题。已知外源施用茉莉酸甲酯(MeJA)或茉莉酸(JA)强烈诱导水稻、小麦、高粱和黑麦草等多种禾本科植物颖花的开放,但是内源JA是否参与水稻颖花开放的调控目前还不清楚。为揭示内源JA在水稻颖花开放中的作用,本研究应用液相色谱-串联质谱系统(HPLC-MS/MS)和实时定量RT-PCR技术(Real-time RT-PCR),检测了粳稻武运粳7号和籼稻明恢63颖花开放前后JA水平及其相关基因表达的变化,并通过水稻转基因方法,探讨了JA生物合成基因OsAOs1和OsOPR7的超量表达或RNAi抑制表达对水稻颖花开放的影响。主要研究结果如下：
(1)水稻成熟颖花JA水平在上午开颖前较平稳,开颖时急剧上升至峰值,闭颖后随即下降。颖花开放时的JA水平比开放前1h提高了4～5倍。
(2)颖花开放过程中JA水平的变化进一步得到了分子基础的支持,JA生物合成途径关键催化酶的编码基因OsDAD1-3、OsLOX-RCI1、OsAOS1、OssAOC和OsOPR7的表达在开颖时会有不同程度的上调,闭颖后又下调。
(3)在阴雨天不开花的条件下,成熟颖花当天JA水平及其合成基因的表达较平稳,不会有峰的出现。
(4)同一稻穗上正在开放的颖花JA水平及其合成基因的表达也比已开花和未开花的颖花高。
(5)通过检测开花前后颖花内各器官JA生物合成基因的表达变化,发现颖花开放过程中大幅增加的JA主要来源于雄蕊和浆片的合成。
(6)JA信号转导途径相关基因OsJAR1、OsCOI1的表达水平在颖花开放过程不随JA生物合成的增加而上升,推测颖花开放过程中JA信号的调控位点不发生在JA信号转导途径上。
(7)使用农杆菌介导的转化方法,获得了JA生物合成基因OsAOS1超量表达(AOS1OE)和RNAi抑制表达(AOS1Ri)转基因植株、以及OsOPR7超量表达(OPR70E)和RNAi抑制表达(OPR7Ri)转基因植株。经表达量检测证实,AOS1OE、OPR70E植株叶片和颖花器官中oSAOS1、OsOPR7的表达水平显著提高,AOS1Ri、OPR7Ri植株叶片和颖花器官中OsAOS1、OSOPR7的表达水平大幅下降。
(8)田间条件下AOS1OE、AOS1Ri、OPR7OE、OPR7Ri植株的颖花器官发育正常,颖花开放时间与野生型对照没有差异,它们的开花高峰时间都在上午11：50左右。
(9)AOS1OE和OPR7OE两个超表达株系的颖花JA水平在开颖前和开颖时都与野生型相比无差异。AOS1Ri和OPR7Ri两个抑制表达株系的颖花JA水平在开颖前与野生型相比也无差异,虽然开颖时不及野生型的50%,但比自身开颖前已有显著增加。结果表明,超量表达OsAOS1和OsOPR7没有增加植株体内基础JA水平,RNAi抑制OsAOS1和OsOPR7的表达也只是部分阻碍了植株体内JA的生物合成,没有达到有效调控植株体内JA水平的目的,这也就合理解释了转基因植株颖花开放时间为什么与野生型无差异。
本研究首次揭示水稻成熟颖花JA水平及其生物合成相关基因表达的变化与颖花开放过程相关,从而为内源JA参与水稻颖花开放的调控提供了有力的证据。
Floret opening of rice which includes opening of palea and lemma, filament elongation and anther dehiscence is a highly organized developmental process, and is controlled by genetic factors and environmental factors (weather conditions and artificial treatments). The regulation of floret opening can contribute to avoid high temperature-induced floret sterility at anthesis, and obtain the synchronization of anthesis between some cytoplasmic male sterile lines and restorer lines in hybrid seed production. It was well documented that exogenously applied jasmonic acid (JA) and methyl jasmonate (MeJA) significantly induce floret opening of rice and other gramineous plants such as wheat, sorghum and ryegrass. However, whether the endogenous JA is involved in the regulation of rice floret opening has remained unclear. To elucidate the role of endogenous JA in rice floret opening, we measured JA levels with HPLC-electrospray ionization tandem mass spectrometry system (HPLC-MS/MS), and analyzed the expression of JA pathway related genes with real-time reverse transcription-PCR (Real-time RT-PCR) in florets of Wuyunjing7(japonica cultivar) and Minghui63(indica cultivar) during natural anthesis. Four transgenic rice lines with overexpression or RNAi-suppression of JA biosynthetic genes OsAOSl or OsOPR7were also generated and analyzed. The main results are as follows:
(1) JA levels in mature florets kept relatively constant before floret opening in the morning, and rapidly reached a peak during floret opening, and then followed a sharp decrease after the floret closing. The peak level of JA in opening florets was4-5times higher than that in florets1h before anthesis.
(2) The changes of JA levels in rice florets during anthesis were supported by our molecular findings. The expression of OsDADl-3, OsLOX-RCIl, OsAOSl, OsAOC and OsOPR7that encode key enzymes in JA biosynthetic pathway was up-regulated to some extent during floret opening, and was down-regulated after floret closing.
(3) The levels of JA and expression of JA biosynthetic genes in unopened mature florets under the condition of overcast did not show obvious changes in the daytime and didn't appear a peak.
(4) The levels of JA and transcripts of JA biosynthetic genes were higher in opening florets than in closed or unopened florets at the same panicle.
(5) The expression of JA related biosynthesis genes was also detected in floret organs before and after anthesis. The results showed that the significantly accumulated JA in opening florets was mainly originated from JA made in stamens and lodicules.
(6) The expression levels of OsJARland OsCOI1involved in JA signal transduction pathway kept rather constant during floret opening, suggesting that JA signal transduction pathway may not be a controlling site for JA pathway during floret opening.
(7) Using the Agrobacterium-mediated rice transformation, we generated four transgenic rice lines, including OsAOS1over-expressed line (AOS1OE), OsAOS1RNAi-suppressed line (AOS1Ri), OsOPR7over-expressed line (OPR7OE) and OsOPR7RNAi-suppressed line (OPR7Ri). mRNA expression level analysis confirmed that OsAOSl transcripts in leaves and florets of AOS1OE lines and OsOPR7transcripts in OPR7OE lines significantly increased, and that OsAOS1transcripts in leaves and florets of AOS1Ri lines and OsOPR7transcripts in OPR7Ri lines significantly reduced.
(8) No obvious difference of spikelet development and floret opening time between the four transgenic lines (AOS1OE, AOS1Ri, OPR7OE and OPR7Ri) and wild-type line was observed in the field condition. The peak time of floret opening in these transgenic and wild-type lines was about at11:50in the morning.
(9) The JA levels in florets of two over-expressed transgenic lines (AOS1OE and OPR7OE) had no significant difference with those of wild-type lines before floret opening and at the time of floret opening. The JA levels in florets of two RNAi-suppressed transgenic lines (AOSIRi and OPR7Ri) were in the same ranges as those of wild-type line before floret opening, and were below half of wild-type line at the time of floret opening. However, the JA levels in RNAi florets at the time of floret opening were significantly increased than that before floret opening. These results indicated that overexpression of OsAOS1and OsOPR7did not change the basal JA level of transgenic plants, and RNAi suppression of OsAOS1and OsOPR7just partially inhibited JA biosynthesis in transgenic plants. This provided a reasonable explanation for that no obvious alteration of floret opening time was observed in the four different transgenic lines in the field condition.
Our study first reported that the changes of JA levels and expression of JA biosynthetic genes in mature florets were associated with process of floret opening, and this strongly indicated that endogenous JA is involved in the regulation of floret opening in rice.
(1)水稻成熟颖花JA水平在上午开颖前较平稳,开颖时急剧上升至峰值,闭颖后随即下降。颖花开放时的JA水平比开放前1h提高了4～5倍。
(2)颖花开放过程中JA水平的变化进一步得到了分子基础的支持,JA生物合成途径关键催化酶的编码基因OsDAD1-3、OsLOX-RCI1、OsAOS1、OssAOC和OsOPR7的表达在开颖时会有不同程度的上调,闭颖后又下调。
(3)在阴雨天不开花的条件下,成熟颖花当天JA水平及其合成基因的表达较平稳,不会有峰的出现。
(4)同一稻穗上正在开放的颖花JA水平及其合成基因的表达也比已开花和未开花的颖花高。
(5)通过检测开花前后颖花内各器官JA生物合成基因的表达变化,发现颖花开放过程中大幅增加的JA主要来源于雄蕊和浆片的合成。
(6)JA信号转导途径相关基因OsJAR1、OsCOI1的表达水平在颖花开放过程不随JA生物合成的增加而上升,推测颖花开放过程中JA信号的调控位点不发生在JA信号转导途径上。
(7)使用农杆菌介导的转化方法,获得了JA生物合成基因OsAOS1超量表达(AOS1OE)和RNAi抑制表达(AOS1Ri)转基因植株、以及OsOPR7超量表达(OPR70E)和RNAi抑制表达(OPR7Ri)转基因植株。经表达量检测证实,AOS1OE、OPR70E植株叶片和颖花器官中oSAOS1、OsOPR7的表达水平显著提高,AOS1Ri、OPR7Ri植株叶片和颖花器官中OsAOS1、OSOPR7的表达水平大幅下降。
(8)田间条件下AOS1OE、AOS1Ri、OPR7OE、OPR7Ri植株的颖花器官发育正常,颖花开放时间与野生型对照没有差异,它们的开花高峰时间都在上午11：50左右。
(9)AOS1OE和OPR7OE两个超表达株系的颖花JA水平在开颖前和开颖时都与野生型相比无差异。AOS1Ri和OPR7Ri两个抑制表达株系的颖花JA水平在开颖前与野生型相比也无差异,虽然开颖时不及野生型的50%,但比自身开颖前已有显著增加。结果表明,超量表达OsAOS1和OsOPR7没有增加植株体内基础JA水平,RNAi抑制OsAOS1和OsOPR7的表达也只是部分阻碍了植株体内JA的生物合成,没有达到有效调控植株体内JA水平的目的,这也就合理解释了转基因植株颖花开放时间为什么与野生型无差异。
本研究首次揭示水稻成熟颖花JA水平及其生物合成相关基因表达的变化与颖花开放过程相关,从而为内源JA参与水稻颖花开放的调控提供了有力的证据。
Floret opening of rice which includes opening of palea and lemma, filament elongation and anther dehiscence is a highly organized developmental process, and is controlled by genetic factors and environmental factors (weather conditions and artificial treatments). The regulation of floret opening can contribute to avoid high temperature-induced floret sterility at anthesis, and obtain the synchronization of anthesis between some cytoplasmic male sterile lines and restorer lines in hybrid seed production. It was well documented that exogenously applied jasmonic acid (JA) and methyl jasmonate (MeJA) significantly induce floret opening of rice and other gramineous plants such as wheat, sorghum and ryegrass. However, whether the endogenous JA is involved in the regulation of rice floret opening has remained unclear. To elucidate the role of endogenous JA in rice floret opening, we measured JA levels with HPLC-electrospray ionization tandem mass spectrometry system (HPLC-MS/MS), and analyzed the expression of JA pathway related genes with real-time reverse transcription-PCR (Real-time RT-PCR) in florets of Wuyunjing7(japonica cultivar) and Minghui63(indica cultivar) during natural anthesis. Four transgenic rice lines with overexpression or RNAi-suppression of JA biosynthetic genes OsAOSl or OsOPR7were also generated and analyzed. The main results are as follows:
(1) JA levels in mature florets kept relatively constant before floret opening in the morning, and rapidly reached a peak during floret opening, and then followed a sharp decrease after the floret closing. The peak level of JA in opening florets was4-5times higher than that in florets1h before anthesis.
(2) The changes of JA levels in rice florets during anthesis were supported by our molecular findings. The expression of OsDADl-3, OsLOX-RCIl, OsAOSl, OsAOC and OsOPR7that encode key enzymes in JA biosynthetic pathway was up-regulated to some extent during floret opening, and was down-regulated after floret closing.
(3) The levels of JA and expression of JA biosynthetic genes in unopened mature florets under the condition of overcast did not show obvious changes in the daytime and didn't appear a peak.
(4) The levels of JA and transcripts of JA biosynthetic genes were higher in opening florets than in closed or unopened florets at the same panicle.
(5) The expression of JA related biosynthesis genes was also detected in floret organs before and after anthesis. The results showed that the significantly accumulated JA in opening florets was mainly originated from JA made in stamens and lodicules.
(6) The expression levels of OsJARland OsCOI1involved in JA signal transduction pathway kept rather constant during floret opening, suggesting that JA signal transduction pathway may not be a controlling site for JA pathway during floret opening.
(7) Using the Agrobacterium-mediated rice transformation, we generated four transgenic rice lines, including OsAOS1over-expressed line (AOS1OE), OsAOS1RNAi-suppressed line (AOS1Ri), OsOPR7over-expressed line (OPR7OE) and OsOPR7RNAi-suppressed line (OPR7Ri). mRNA expression level analysis confirmed that OsAOSl transcripts in leaves and florets of AOS1OE lines and OsOPR7transcripts in OPR7OE lines significantly increased, and that OsAOS1transcripts in leaves and florets of AOS1Ri lines and OsOPR7transcripts in OPR7Ri lines significantly reduced.
(8) No obvious difference of spikelet development and floret opening time between the four transgenic lines (AOS1OE, AOS1Ri, OPR7OE and OPR7Ri) and wild-type line was observed in the field condition. The peak time of floret opening in these transgenic and wild-type lines was about at11:50in the morning.
(9) The JA levels in florets of two over-expressed transgenic lines (AOS1OE and OPR7OE) had no significant difference with those of wild-type lines before floret opening and at the time of floret opening. The JA levels in florets of two RNAi-suppressed transgenic lines (AOSIRi and OPR7Ri) were in the same ranges as those of wild-type line before floret opening, and were below half of wild-type line at the time of floret opening. However, the JA levels in RNAi florets at the time of floret opening were significantly increased than that before floret opening. These results indicated that overexpression of OsAOS1and OsOPR7did not change the basal JA level of transgenic plants, and RNAi suppression of OsAOS1and OsOPR7just partially inhibited JA biosynthesis in transgenic plants. This provided a reasonable explanation for that no obvious alteration of floret opening time was observed in the four different transgenic lines in the field condition.
Our study first reported that the changes of JA levels and expression of JA biosynthetic genes in mature florets were associated with process of floret opening, and this strongly indicated that endogenous JA is involved in the regulation of floret opening in rice.
引文
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