小麦抗逆相关转录因子bZIP和NAC基因的功能研究
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摘要
bZIP和NAC是植物中两个重要的转录因子家族，它们参与调控植物的生长发育及逆境胁迫应答等过程。抗逆相关的bZIP和NAC转录因子的功能在拟南芥和水稻中研究较为深入，而在异源六倍体小麦中的研究相对较少。我们在小麦中对这两个家族的基因进行了研究，期望发掘一些与抗逆相关的小麦bZIP和NAC转录因子基因，为小麦的分子育种提供候选基因。我们从本实验室构建的小麦全长cDNA文库中，筛选了22个TabZIP和28个TaNAC基因。通过半定量RT-PCR方法，对这些基因的组织特异性及响应非生物胁迫的表达模式进行了研究，筛选了一些与非生物胁迫相关的TabZIP和TaNAC基因。并在拟南芥中进行了过表达研究，具体结果如下：
1、组织特异性表达分析结果表明，9个TabZIP和18个TaNAC基因在幼穗、叶、茎和根中都表达，13个TabZIP和10个TaNAC基因在不同组织中呈现不同水平的表达。低温胁迫条件下，10个TabZIP和13个TaNAC基因表达量上调；13个TabZIP和15个TaNAC基因受盐胁迫诱导表达；在PEG诱导的渗透胁迫下，10个TabZIP基因表达量上调；15个TabZIP基因表达受ABA上调。
2、TabZIP1属于bZIP转录因子A家族的基因，定位于小麦第六同源群的6A、6B和6D上。该基因编码一个定位在细胞核中的蛋白，在酵母中具有转录激活活性。TabZIP1受盐、PEG、低温和外源ABA诱导表达，对过表达TabZIP1的转基因拟南芥进行抗逆性鉴定表明：TabZIP1提高了转基因拟南芥对盐、干旱和冻害的抗性。酵母单杂交结果表明该基因与ABRE顺式元件相互作用。并且过表达TabZIP1的转基因拟南芥增加了对ABA的敏感性，进一步表明TabZIP1参与调控ABA信号途径。
3、TabZIP14属于bZIP转录因子C家族的基因，编码一个定位在细胞核中的蛋白，在酵母中具有转录激活活性。TabZIP14基因受盐、PEG、低温和外源ABA诱导表达，对过表达TabZIP14的转基因拟南芥进行抗逆性鉴定表明：TabZIP14提高了转基因拟南芥对盐和冻害的抗性。
4、TaNAC47属于NAC转录因子SNAC家族的基因，编码一个定位在细胞核中的蛋白，在酵母中具有转录激活活性。TaNAC47基因受多种逆境胁迫的诱导表达，对过表达TaNAC47的转基因拟南芥进行抗逆性鉴定表明：TaNAC47提高了转基因拟南芥对盐、干旱和冻害的抗性。
5、TaNAC62属于NAC转录因子NTLs家族的基因，定位于小麦第三同源群的3A、3B和3D上。该基因编码一个定位在细胞核中的蛋白，在酵母中具有转录激活活性。TaNAC62基因受多种逆境胁迫的诱导表达，对过表达TaNAC62的转基因拟南芥进行抗逆性鉴定表明：TaNAC62提高了转基因拟南芥对盐、干旱和冻害的抗性。
The basic region/leucine zipper (bZIP) and NAC that are pivotal regulators involved ingrowth, development of plants and abiotic stress are important transcription factors in plants. InArabidopsis and rice, many bZIP genes have been reported with roles in response to abioticstresses. However, there are few reports on the stress-related bZIP genes in allohexaploid wheat.In this study, we conducted research concerning the function of wheat transcription factor bZIPand NAC involved in abiotic stress tolerances, which wound provide candidate genes for wheatmolecular breeding in the future. The22TabZIP and28TaNAC genes were screened from wheatfull-length cDNA libraries in our laboratory. The expression patterns of the tissue-specific andabiotic stress response of these genes were studied by semi-QRT-PCR. Some stress-related TabZIPand TaNAC genes were obtained. The functions of these stress-related TabZIP and TaNAC geneswere carried out through ectopic overexpression in Arabidopsis. The main results are as follows:
1. Tissue specific anlysis indicated9TabZIP and18TaNAC were expressed in young spikes,leaves, stems and roots. The13TabZIP and10TaNAC genes were expressed differently indifferent tissues. The expression levels of10TabZIP and13TaNAC genes were up-regulatedunder low temperature stress. The expression of13TabZIP and15TaNAC genes increased undersalt stress treatment. For polyethylene glycol (PEG)-induced osmotic stress, the transcriptionexpression of10TabZIP genes enhanced. The expression of15TabZIP genes were induced byexogenous abscisic acid (ABA) treatment.
2. TabZIP1gene belonged to the A family of bZIP transcription factor, which was mapped tothe wheat chromosomes6A,6B and6D. TabZIP1protein was localized to the nucleus and hadtranscriptional activity in yeast. TabZIP1gene transcripts were strongly induced by salt, PEG, lowtemperature and exogenous ABA treatments. Over-expression of TabZIP1in Arabidopsis led toenhanced tolerances to salt, drought and freezing stresses. The TabZIP1was capable of bindingABRE sequence by yeast one-hybrid assay and the overexpression of TabZIP1in Arabidopsis ledto greater sensitivity to exogenous ABA in seedling growth. The results indicated TabZIP1participated in ABA signaling pathway.
3. TabZIP14gene belonged to the C family of bZIP transcription factor. TabZIP14proteinwas localized to the nucleus and had transcriptional activity in yeast. TabZIP14gene transcriptswere strongly induced by salt, PEG, low temperature and ABA treatments. Over-expression ofTabZIP14in Arabidopsis led to enhanced tolerances to salt and freezing stresses.
4. TaNAC47gene belonged to SNAC family of NAC transcription factor. TaNAC47proteinwas localized to the nucleus and had transcriptional activity in yeast. TaNAC47gene transcriptswere strongly induced by salt, PEG, low temperature and ABA treatments. Over-expression of TaNAC47in Arabidopsis led to enhanced tolerances to salt, drought and freezing stresses.
5. TaNAC62gene belonged to NTLs family of NAC transcription factor, which was mappedto the wheat chromosomes3A,3B and3D. TaNAC62protein was localized to the nucleus andhad transcriptional activity in yeast. TaNAC62gene transcripts were strongly induced by salt,PEG, low temperature and ABA treatments. Over-expression of TaNAC62in Arabidopsis led toenhanced tolerances to salt, drought and freezing stresses.
1、组织特异性表达分析结果表明，9个TabZIP和18个TaNAC基因在幼穗、叶、茎和根中都表达，13个TabZIP和10个TaNAC基因在不同组织中呈现不同水平的表达。低温胁迫条件下，10个TabZIP和13个TaNAC基因表达量上调；13个TabZIP和15个TaNAC基因受盐胁迫诱导表达；在PEG诱导的渗透胁迫下，10个TabZIP基因表达量上调；15个TabZIP基因表达受ABA上调。
2、TabZIP1属于bZIP转录因子A家族的基因，定位于小麦第六同源群的6A、6B和6D上。该基因编码一个定位在细胞核中的蛋白，在酵母中具有转录激活活性。TabZIP1受盐、PEG、低温和外源ABA诱导表达，对过表达TabZIP1的转基因拟南芥进行抗逆性鉴定表明：TabZIP1提高了转基因拟南芥对盐、干旱和冻害的抗性。酵母单杂交结果表明该基因与ABRE顺式元件相互作用。并且过表达TabZIP1的转基因拟南芥增加了对ABA的敏感性，进一步表明TabZIP1参与调控ABA信号途径。
3、TabZIP14属于bZIP转录因子C家族的基因，编码一个定位在细胞核中的蛋白，在酵母中具有转录激活活性。TabZIP14基因受盐、PEG、低温和外源ABA诱导表达，对过表达TabZIP14的转基因拟南芥进行抗逆性鉴定表明：TabZIP14提高了转基因拟南芥对盐和冻害的抗性。
4、TaNAC47属于NAC转录因子SNAC家族的基因，编码一个定位在细胞核中的蛋白，在酵母中具有转录激活活性。TaNAC47基因受多种逆境胁迫的诱导表达，对过表达TaNAC47的转基因拟南芥进行抗逆性鉴定表明：TaNAC47提高了转基因拟南芥对盐、干旱和冻害的抗性。
5、TaNAC62属于NAC转录因子NTLs家族的基因，定位于小麦第三同源群的3A、3B和3D上。该基因编码一个定位在细胞核中的蛋白，在酵母中具有转录激活活性。TaNAC62基因受多种逆境胁迫的诱导表达，对过表达TaNAC62的转基因拟南芥进行抗逆性鉴定表明：TaNAC62提高了转基因拟南芥对盐、干旱和冻害的抗性。
The basic region/leucine zipper (bZIP) and NAC that are pivotal regulators involved ingrowth, development of plants and abiotic stress are important transcription factors in plants. InArabidopsis and rice, many bZIP genes have been reported with roles in response to abioticstresses. However, there are few reports on the stress-related bZIP genes in allohexaploid wheat.In this study, we conducted research concerning the function of wheat transcription factor bZIPand NAC involved in abiotic stress tolerances, which wound provide candidate genes for wheatmolecular breeding in the future. The22TabZIP and28TaNAC genes were screened from wheatfull-length cDNA libraries in our laboratory. The expression patterns of the tissue-specific andabiotic stress response of these genes were studied by semi-QRT-PCR. Some stress-related TabZIPand TaNAC genes were obtained. The functions of these stress-related TabZIP and TaNAC geneswere carried out through ectopic overexpression in Arabidopsis. The main results are as follows:
1. Tissue specific anlysis indicated9TabZIP and18TaNAC were expressed in young spikes,leaves, stems and roots. The13TabZIP and10TaNAC genes were expressed differently indifferent tissues. The expression levels of10TabZIP and13TaNAC genes were up-regulatedunder low temperature stress. The expression of13TabZIP and15TaNAC genes increased undersalt stress treatment. For polyethylene glycol (PEG)-induced osmotic stress, the transcriptionexpression of10TabZIP genes enhanced. The expression of15TabZIP genes were induced byexogenous abscisic acid (ABA) treatment.
2. TabZIP1gene belonged to the A family of bZIP transcription factor, which was mapped tothe wheat chromosomes6A,6B and6D. TabZIP1protein was localized to the nucleus and hadtranscriptional activity in yeast. TabZIP1gene transcripts were strongly induced by salt, PEG, lowtemperature and exogenous ABA treatments. Over-expression of TabZIP1in Arabidopsis led toenhanced tolerances to salt, drought and freezing stresses. The TabZIP1was capable of bindingABRE sequence by yeast one-hybrid assay and the overexpression of TabZIP1in Arabidopsis ledto greater sensitivity to exogenous ABA in seedling growth. The results indicated TabZIP1participated in ABA signaling pathway.
3. TabZIP14gene belonged to the C family of bZIP transcription factor. TabZIP14proteinwas localized to the nucleus and had transcriptional activity in yeast. TabZIP14gene transcriptswere strongly induced by salt, PEG, low temperature and ABA treatments. Over-expression ofTabZIP14in Arabidopsis led to enhanced tolerances to salt and freezing stresses.
4. TaNAC47gene belonged to SNAC family of NAC transcription factor. TaNAC47proteinwas localized to the nucleus and had transcriptional activity in yeast. TaNAC47gene transcriptswere strongly induced by salt, PEG, low temperature and ABA treatments. Over-expression of TaNAC47in Arabidopsis led to enhanced tolerances to salt, drought and freezing stresses.
5. TaNAC62gene belonged to NTLs family of NAC transcription factor, which was mappedto the wheat chromosomes3A,3B and3D. TaNAC62protein was localized to the nucleus andhad transcriptional activity in yeast. TaNAC62gene transcripts were strongly induced by salt,PEG, low temperature and ABA treatments. Over-expression of TaNAC62in Arabidopsis led toenhanced tolerances to salt, drought and freezing stresses.
引文
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