转Bcl-2、Ced-9、PpBl-1基因水稻抗盐性及其抗性机理研究
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摘要
盐胁迫是植物生长发育的主要限制因素之一,严重影响着农作物的产量和品质。利用基因工程手段提高农作物的抗盐性已成为作物遗传改良的重要内容之一。前人的研究结果已经证实凋亡抑制基因Bc]-2和Ced-9能够提高植物细胞对生物和非生物胁迫的抗性,但是其作用机制仍知之甚少。本研究以水稻为材料,开展盐(NaCl)诱导细胞程序性死亡(programmed cell death, PCD)的研究,并通过转细胞凋亡抑制基因Bcl-2/Ced-9/PpBI-1提高水稻的耐盐性,研究其耐性机理。获得如下主要结果：
1Bcl-2基因通过抑制盐胁迫引起的胞质Ca2+增加和PCD的产生,提高水稻对盐胁迫的耐受性。
1.1在盐(NaCl)胁迫条件下,Bcl-2转基因水稻比野生型具有较高的萌发率、相对根长生长和细胞活性,显示更强的耐盐性,证明凋亡抑制基因Bcl-2在转基因水稻根尖细胞内具有明显的促进耐盐胁迫功能。
1.2用DNA ladder检测方法,比较转Bcl-2基因和野生型水稻在盐胁迫条件下的细胞PCD,发现Bcl-2基因的过表达能明显地抑制盐诱导PCD的发生,说明过表达Bcl-2转基因水稻通过抑制PCD的产生而提高耐盐胁迫能力。
1.3用Fluo-3/AM共聚焦荧光显微技术,发现NaCl能显著诱导水稻根尖细胞内钙离子(Ca2+)水平增加；Bcl-2能直接抑制调节胞内钙离子(Ca2+)水平。提示Bcl-2作用于盐胁迫引起的Ca2+信号水平的调控。Bcl-2在植物细胞中的耐盐机制可能是通过控制细胞内Ca2+信号水平的升高来抑制盐胁迫引起的PCD。
1.4La3+处理能显著缓解野生型盐胁迫导致的水稻萌发率和根伸长抑制,减缓胞内钙离子(Ca2+)水平上升,抑制PCD发生。但La3+处理对Bcl-2转基因水稻作用不明显。
2用非损伤微测技术(Non-invasive Microtest Technique(NMT))进一步证明了Bcl-2通过调节离子流抑制盐胁迫引起的PCD。
2.1用非损伤微测技术(Non-invasive Microtest Technique),测定盐胁迫引起的水稻根尖细胞的离子流,结果表明,在盐胁迫下,野生型水稻细胞产生明显的K+和H+外流,而Bcl-2誓能够显著减弱K+外流并抑制PCD的产生,但H+外流没有影响。证明在盐胁迫下,发生水稻根尖细胞PCD的主要原因是盐胁迫引起的K+外流。
2.2La3+处理野生型能够抑制盐胁迫引起的水稻根尖细胞K+外流,而对转Bcl-2基因水稻的作用不明显。说明Bcl-2的作用位点可能与La3+作用位点相同,均与NSCC发生作用。
2.3采用共聚焦显微技术,测定细胞内Ca2+的含量以及Ca2+外流,结果表明在盐胁迫下,水稻根尖细胞Ca2+外流增加,而Bcl-2转基因水稻的Ca2+外排能力比野生型更强。
3在盐胁迫-下,野生型水稻OsVPE-2和OsVPE-3表达显著高于Bcl-2转基因水稻；La3+处理能显著抑制野生型盐胁迫引起的OsVPE-2和OsVPE-3表达,但对Bcl-2转基因水稻影响不明显。
3.1定量PCR测定结果表明,与野生型相比,胞质Ca2+水平显著降低的过表达Bc1-2转基因水稻中OsVPE-2和OsVPE-3的表达显著被抑制。
3.2在盐胁迫条件下,La3+处理能显著抑制野生型盐胁迫引起的OsVPE-2和OsVPE-3表达,而La3+处理对转Bcl-2基因水稻OsVPE-2和OsVPE-3表达的影响不明显。
3.3综合K+、Ca2+测定结果和OsVPE-2和OsVPE-3表达特征,表明在盐胁迫条件下,通过改变K+外流,影响胞质K+、Ca2+浓度,调节OsVPE-2和OsVPE-3表达,影响PCD的产生,最后表现出不同的抗盐性。
4通过根癌农杆菌(Agrobacterium tumefaciens) EHA105介导转化方法,成功将抗凋亡基因Ced-9和PpBI-1转入水稻细胞,获得了转Ced-9和PpBI-1基因水稻株系,并进行抗盐性鉴定及其遗传分析。
4.1构建含Ced-9或PpBI-1基因的表达质粒pCAMBIA13011-Ced-9/PpBI-1,通过根癌农杆菌EHA105介导转化水稻(日本晴),经潮霉素筛选和PCR鉴定、Southern杂交分析,证明Ced-9和PpBI-1基因成功插入到水稻基因组,并用RT-PCR分析确认Ced-9和PpBI-11基因已成功表达,获得了两个转基因纯合株系。
4.2通过对转Ced-9/PpBI-1基因水稻T3代的生长发育特性分析,观察到转Ced-9基因水稻生长发育正常,而转PpBI-1基因水稻生长发育受到一定影响。认为与转基因插入位点有关。
4.3通过对转Ced-9/PpBI-1基因水稻T3代的抗盐性鉴定,发现转Ced-9/PpBI-1基因水稻株系在萌发率和根伸长方面明显表现出对盐胁迫的耐受性。
4.4通过DNA laddering检测发现,Ced-9基因过表达能减弱盐胁迫引起的水稻根尖细胞PCD。
NaCl stress is the major factor limiting plant growth and development, which seriously affects the yields and qualities of crops. Using transgenic technique has been an important strategy to produce agricultural crops to improve stress resistance. It has been shown that apoptotic suppressors such as Bcl-xL and Ced-9can enhance the resistance of plant cells to biotic or abiotic stresses, but detailed mechanisms remain still largely unclear. Here, we choose two rice strains(Oryza sativa L.subsp.Japonica cultivar Zhonghuall; Oryza sativa cv. Nipponbare) as an excellent model for studying various aspects of salt induced programmed cell death (PCD). And our studies were carried out to elucidate the mechanisms and enhance the resistant to salt stress by overexpressing Bcl-2/Ced-9/PpB1-1in rice. Results obtaind in our research are summarized as follows:
1Bcl-2improves the tolerance to salt stress via suppressing cytosolic Ca2+increase and PCD caused by salt stress in rice.
1.1Bcl-2transgenic rice possessed the ability to overcome NaCl stress. Under saline conditions, it can obviously improve the seed germination, root elongation and cell viability.
1.2The profile of DNA laddering indicates that NaCl induces cell death accompanied by typical hallmarks of PCD, while Bcl-2could promote NaCl tolerance in transgenic rice via inhibition of PCD.
1.3Investigation with Fluo-3/AM CONFOCAL shows that salt stress lead to increase cytosolic Ca2+in WT line, contrast to in Bcl-2line, suggesting that Bcl-2mediate closely with Ca2+signaling pathway induced upon salt stress.
1.4La3+has the distinct ability to improve the seed germination and root elongation, reduce cytosolic Ca2+and inhibit DNA laddering in WT line, but not in Bcl-2line.
2Using the Non-invasive Micro-test Technique (NMT), we confirm that under salt stress, Bcl-2can suppress NaCI-induced PCD by means of the regulation of ion fluxesin in rice cell.
2.1The main cellular events under salt stress involve K+efflux from rice tip cells, which may be effectively suppressed by Bcl-2, so that salt-induced PCD can be inhibited. Otherwise, H+efflux does not change in such condition.
2.2La3+can contribute to inhibition of K+efflux under salt stress in WT rice. However, its potential indicates unclearly in Bcl-2line.
2.3Ca2+efflux is a common phenomenon under salt stress, which undertakes to much more extent in Bcl-2than in WT line.
3K+efflux in cytoplasm under salt stress lead to transcriptional activation of OsVPE-2and OsVPE-3, furthermore, results in PCD. Such activation can be remarkably down regulated by La3+in WT rice, but not in Bcl-2line.
3.1Q-RT-PCR detection showed that salt stress significantly increase expression of OsVPE-2and OsVPE-3in WT, but the expressions were inhibited in Bcl-2line.
3.2Under salt sress, La3+treatment inhibited expression of OsVPE-2and OsVPE-3in WT, but not in in Bcl-2line.
4Ced-9and PpBI-1gene were tansformed into rice calli by Agrobacterium tumefaciens-mediated transformation. Two transgenic lines, which possessed tolerance to salt, were successfully obtained in rice.
4.1The incorporation of pCAMBIA13011-Ced-9/PpB1-1into rice genome was detected/identified by hygromycin selection, and PCR relative to target genes and Southern blotting. The expression of Ced-9/PpBl-1were confirmed by RT-PCR analysis in transgenic lines, and as a result, homozygotic lines of transgenic rice were obtained in two lines.
4.2As with the investigations of pCAMBIA13011-Ced-9/PpBI-1transgenic T3lines, Ced-9line shows normal developmental phase, but not in PpBI-1line, suggesting that it would be probably caused by the different locus of T-DNA insertion.
4.3Ced-9/PpBI-1transgenic T3lines possessed tolerance to salt, with respect to the seed germination and root elongation.
4.4Through detailed analysis of DNA laddering, Ced-9was recognized as an effective factor to suppress NaCl-induced PCD, in rice.
1Bcl-2基因通过抑制盐胁迫引起的胞质Ca2+增加和PCD的产生,提高水稻对盐胁迫的耐受性。
1.1在盐(NaCl)胁迫条件下,Bcl-2转基因水稻比野生型具有较高的萌发率、相对根长生长和细胞活性,显示更强的耐盐性,证明凋亡抑制基因Bcl-2在转基因水稻根尖细胞内具有明显的促进耐盐胁迫功能。
1.2用DNA ladder检测方法,比较转Bcl-2基因和野生型水稻在盐胁迫条件下的细胞PCD,发现Bcl-2基因的过表达能明显地抑制盐诱导PCD的发生,说明过表达Bcl-2转基因水稻通过抑制PCD的产生而提高耐盐胁迫能力。
1.3用Fluo-3/AM共聚焦荧光显微技术,发现NaCl能显著诱导水稻根尖细胞内钙离子(Ca2+)水平增加；Bcl-2能直接抑制调节胞内钙离子(Ca2+)水平。提示Bcl-2作用于盐胁迫引起的Ca2+信号水平的调控。Bcl-2在植物细胞中的耐盐机制可能是通过控制细胞内Ca2+信号水平的升高来抑制盐胁迫引起的PCD。
1.4La3+处理能显著缓解野生型盐胁迫导致的水稻萌发率和根伸长抑制,减缓胞内钙离子(Ca2+)水平上升,抑制PCD发生。但La3+处理对Bcl-2转基因水稻作用不明显。
2用非损伤微测技术(Non-invasive Microtest Technique(NMT))进一步证明了Bcl-2通过调节离子流抑制盐胁迫引起的PCD。
2.1用非损伤微测技术(Non-invasive Microtest Technique),测定盐胁迫引起的水稻根尖细胞的离子流,结果表明,在盐胁迫下,野生型水稻细胞产生明显的K+和H+外流,而Bcl-2誓能够显著减弱K+外流并抑制PCD的产生,但H+外流没有影响。证明在盐胁迫下,发生水稻根尖细胞PCD的主要原因是盐胁迫引起的K+外流。
2.2La3+处理野生型能够抑制盐胁迫引起的水稻根尖细胞K+外流,而对转Bcl-2基因水稻的作用不明显。说明Bcl-2的作用位点可能与La3+作用位点相同,均与NSCC发生作用。
2.3采用共聚焦显微技术,测定细胞内Ca2+的含量以及Ca2+外流,结果表明在盐胁迫下,水稻根尖细胞Ca2+外流增加,而Bcl-2转基因水稻的Ca2+外排能力比野生型更强。
3在盐胁迫-下,野生型水稻OsVPE-2和OsVPE-3表达显著高于Bcl-2转基因水稻；La3+处理能显著抑制野生型盐胁迫引起的OsVPE-2和OsVPE-3表达,但对Bcl-2转基因水稻影响不明显。
3.1定量PCR测定结果表明,与野生型相比,胞质Ca2+水平显著降低的过表达Bc1-2转基因水稻中OsVPE-2和OsVPE-3的表达显著被抑制。
3.2在盐胁迫条件下,La3+处理能显著抑制野生型盐胁迫引起的OsVPE-2和OsVPE-3表达,而La3+处理对转Bcl-2基因水稻OsVPE-2和OsVPE-3表达的影响不明显。
3.3综合K+、Ca2+测定结果和OsVPE-2和OsVPE-3表达特征,表明在盐胁迫条件下,通过改变K+外流,影响胞质K+、Ca2+浓度,调节OsVPE-2和OsVPE-3表达,影响PCD的产生,最后表现出不同的抗盐性。
4通过根癌农杆菌(Agrobacterium tumefaciens) EHA105介导转化方法,成功将抗凋亡基因Ced-9和PpBI-1转入水稻细胞,获得了转Ced-9和PpBI-1基因水稻株系,并进行抗盐性鉴定及其遗传分析。
4.1构建含Ced-9或PpBI-1基因的表达质粒pCAMBIA13011-Ced-9/PpBI-1,通过根癌农杆菌EHA105介导转化水稻(日本晴),经潮霉素筛选和PCR鉴定、Southern杂交分析,证明Ced-9和PpBI-1基因成功插入到水稻基因组,并用RT-PCR分析确认Ced-9和PpBI-11基因已成功表达,获得了两个转基因纯合株系。
4.2通过对转Ced-9/PpBI-1基因水稻T3代的生长发育特性分析,观察到转Ced-9基因水稻生长发育正常,而转PpBI-1基因水稻生长发育受到一定影响。认为与转基因插入位点有关。
4.3通过对转Ced-9/PpBI-1基因水稻T3代的抗盐性鉴定,发现转Ced-9/PpBI-1基因水稻株系在萌发率和根伸长方面明显表现出对盐胁迫的耐受性。
4.4通过DNA laddering检测发现,Ced-9基因过表达能减弱盐胁迫引起的水稻根尖细胞PCD。
NaCl stress is the major factor limiting plant growth and development, which seriously affects the yields and qualities of crops. Using transgenic technique has been an important strategy to produce agricultural crops to improve stress resistance. It has been shown that apoptotic suppressors such as Bcl-xL and Ced-9can enhance the resistance of plant cells to biotic or abiotic stresses, but detailed mechanisms remain still largely unclear. Here, we choose two rice strains(Oryza sativa L.subsp.Japonica cultivar Zhonghuall; Oryza sativa cv. Nipponbare) as an excellent model for studying various aspects of salt induced programmed cell death (PCD). And our studies were carried out to elucidate the mechanisms and enhance the resistant to salt stress by overexpressing Bcl-2/Ced-9/PpB1-1in rice. Results obtaind in our research are summarized as follows:
1Bcl-2improves the tolerance to salt stress via suppressing cytosolic Ca2+increase and PCD caused by salt stress in rice.
1.1Bcl-2transgenic rice possessed the ability to overcome NaCl stress. Under saline conditions, it can obviously improve the seed germination, root elongation and cell viability.
1.2The profile of DNA laddering indicates that NaCl induces cell death accompanied by typical hallmarks of PCD, while Bcl-2could promote NaCl tolerance in transgenic rice via inhibition of PCD.
1.3Investigation with Fluo-3/AM CONFOCAL shows that salt stress lead to increase cytosolic Ca2+in WT line, contrast to in Bcl-2line, suggesting that Bcl-2mediate closely with Ca2+signaling pathway induced upon salt stress.
1.4La3+has the distinct ability to improve the seed germination and root elongation, reduce cytosolic Ca2+and inhibit DNA laddering in WT line, but not in Bcl-2line.
2Using the Non-invasive Micro-test Technique (NMT), we confirm that under salt stress, Bcl-2can suppress NaCI-induced PCD by means of the regulation of ion fluxesin in rice cell.
2.1The main cellular events under salt stress involve K+efflux from rice tip cells, which may be effectively suppressed by Bcl-2, so that salt-induced PCD can be inhibited. Otherwise, H+efflux does not change in such condition.
2.2La3+can contribute to inhibition of K+efflux under salt stress in WT rice. However, its potential indicates unclearly in Bcl-2line.
2.3Ca2+efflux is a common phenomenon under salt stress, which undertakes to much more extent in Bcl-2than in WT line.
3K+efflux in cytoplasm under salt stress lead to transcriptional activation of OsVPE-2and OsVPE-3, furthermore, results in PCD. Such activation can be remarkably down regulated by La3+in WT rice, but not in Bcl-2line.
3.1Q-RT-PCR detection showed that salt stress significantly increase expression of OsVPE-2and OsVPE-3in WT, but the expressions were inhibited in Bcl-2line.
3.2Under salt sress, La3+treatment inhibited expression of OsVPE-2and OsVPE-3in WT, but not in in Bcl-2line.
4Ced-9and PpBI-1gene were tansformed into rice calli by Agrobacterium tumefaciens-mediated transformation. Two transgenic lines, which possessed tolerance to salt, were successfully obtained in rice.
4.1The incorporation of pCAMBIA13011-Ced-9/PpB1-1into rice genome was detected/identified by hygromycin selection, and PCR relative to target genes and Southern blotting. The expression of Ced-9/PpBl-1were confirmed by RT-PCR analysis in transgenic lines, and as a result, homozygotic lines of transgenic rice were obtained in two lines.
4.2As with the investigations of pCAMBIA13011-Ced-9/PpBI-1transgenic T3lines, Ced-9line shows normal developmental phase, but not in PpBI-1line, suggesting that it would be probably caused by the different locus of T-DNA insertion.
4.3Ced-9/PpBI-1transgenic T3lines possessed tolerance to salt, with respect to the seed germination and root elongation.
4.4Through detailed analysis of DNA laddering, Ced-9was recognized as an effective factor to suppress NaCl-induced PCD, in rice.
引文
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Lawen A.2003. Apoptosis-an introduction. Bioessays 25:888-896.
Letai A.2005. Pharmacological manipulation of Bcl-2 family members to control cell death. J Clin Invest 115:2648-2655.
Li J-Y, Jiang A-L, Chen H-Y, Wang Y, Zhang W.2007. Lanthanum Prevents Salt Stress-induced Programmed Cell Death in Rice Root Tip Cells by Controlling Early Induction Events. Journal of Integrative Plant Biology 49:1024-1031.
Li X, et al.2011. Regulating cytoplasmic calcium homeostasis can reduce aluminum toxicity in yeast. PLoS One 6:e21148.
Lin J, Wang Y, Wang G. 2005. Salt stress-induced programmed cell death via Ca2+-mediated mitochondrial permeability transition in tobacco protoplasts. Plant Growth Regulation 45:243-250.
Lincoln JE, Richael C, Overduin B, Smith K, Bostock R, Gilchrist DG. 2002. Expression of the antiapoptotic baculovirus p35 gene in tomato blocks programmed cell death and provides broad-spectrum resistance to disease. Proc Natl Acad Sci U S A 99:15217-15221.
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