小麦类钙调磷酸酶B亚基蛋白TaCBL4和TaCBL3的功能研究
摘要
非生物逆境是作物产量的主要制约因素。现代分子生物学手段现已广泛应用于育种研究,并获得不少具有优良耐逆能力的作物新品种。类钙调磷酸酶B亚基蛋白(calcineurin B-like proteins, CBL)是一类重要的钙信号受体蛋白,在非生物胁迫应答反应中具有重要作用,主要通过与CBL相互作用蛋白激酶(CBL-interacting protein kinases, CIPK)相互作用将信号向下游传递。拟南芥中CBL和CIPK家族成员在耐逆中的作用已有较深入的研究,其中AtSOS3(CBL4)和AtCIPK24(SOS2)是SOS途径的两个关键组分,在植物盐胁迫应答中发挥重要作用。但是小麦中CBL和CIPK家族成员功能以及是否存在SOS途径还不清楚,SOS途径在植物中是否具有保守性和通用性还未见报道。
本研究基于以上科学问题,利用生物信息学方法系统分析了小麦CBL和CIPK基因家族,从本室培育的小麦渐渗系耐盐品种山融3号(SR3)中克隆了一系列TaCBLs和TaCIPKs基因,并初步验证了TaCBL4、TaCBL3在非生物胁迫应答中的功能。
一、小麦CBL和CIPK的生物信息学分析和新基因克隆
通过生物信息学分析,电子克隆了14个小麦CBL家族成员和34个小麦CIPK家族成员。多序列比对及进化树分析显示,这些TaCBLs在大小和结构上非常保守,其中5个具有豆蔻酰化位点,2个具有跨膜疏水区,暗示这7个TaCBLs在抗逆中发挥重要作用。TaCIPKs间序列差异比较大,但结构十分保守。SR3中TaCIPKs间也存在差异,这可能与基因倍增过程相关。进化树分析显示,小麦中TaCBLs和TaCIPKs都存在许多旁系同源基因,并且它们与水稻中同源基因的相似性高于拟南芥。
将这些电子克隆的TaCIPK基因与实验室定制的小麦cDNA芯片探针进行比对,以分析它们的盐胁迫应答特征。结果发现,8个TaCIPKs在芯片中有特异性匹配探针,其中4个为盐诱导表达基因,4个盐下调表达基因：另外,芯片中有3个探针分别对应多个TaCIPK序列,而且这3个探针在盐胁迫24h后表达量都显著提高。这些盐胁迫应答的TaCIPKs(?)可能都参与植物的盐胁迫应答反应,其功能需要进一步研究。
根据电子克隆序列,从SR3中克隆3个TaCBL和9个TaCIPK新基因,按照与拟南芥CBLs的同源性,分别命名为TaCBL1/3/4和TaC1PK2/8/11/15/17/19/30/32/34。 TaClPKs中,TaCIPK8、TaCIPK15在盐胁迫下上调表达,TaCIPK11下调表达,其他基因的应答模式有待进一步验证,为耐盐候选基因的鉴定及TaCBL互作蛋白的筛选奠定了基础。
二、小麦TaCBL4和TaCBL3功能分析
1、TaCBL4结构与可能作用分析
TaCBL4有7个内含子,其编码产物定位于细胞膜,与拟南芥同源基因AtSOS3(CBL4)相似,是AtSOS3同源基因。TaCBL4在小麦的根、叶、茎、幼穗、麦芒、幼胚、旗叶等组织中均能表达,NaCl(?)协迫下其表达量明显增加,表明TaCBL4在小麦生长发育和盐胁迫应答过程中均发挥重要作用。
为了验证TaCBL4(?)的生物学功能及其与AtSOS3(?)的异同,将其在拟南芥Col-0野生型和sos3突变体中异源表达,观察表型变化,并用AtSOS3在拟南芥中的过表达系作为对照。TaCBL4过表达不影响拟南芥的营养和生殖生长。在1/2MS培养基中,Col-0(WT)、TaCBL4过表达野生型系(OE)、TaCBL4过表达sos3(sOE)幼苗根长和地上部分无显著性差异。NaCl胁迫下,与WT相比,sos3幼苗的主根更短、地上部分漂白程度高,与朱建康实验室结果一致；而OE系和sOE系幼苗根与WT没有显著差别,但是叶片显著变小。这显示,TaCBL4能够恢复sos3根的表型,但首次发现OE系和sOE系在胁迫下抑制叶生长。LiCl胁迫下,与WT相比,sos3幼苗的主根明显变短、地上部分变小,而OH系、sOE系及AtSOS3过表达系幼苗正好相反,主根更长、地上部分更大,首次表明TaCBL4和AtSOS3过表达可以提高幼苗期锂离子胁迫抗性。在ABA处理和渗透胁迫下不同株系无明显差异,表明TaCBL4是一个离子特异性响应基因。
相比植株发育,盐胁迫对种子萌发的影响更大。为了分析TaCBL4在这阶段是否发挥重要作用,我们以绿色子叶张开率为标准,比较了不同株系在钠、锂离子胁迫下的萌发率差异。NaCl胁迫下,sos3不萌发,WT萌发率降低,而OE系和sOE萌发率受影响少,萌发率明显比WT高。LiCl处理下,WT和sos3的萌发率降低,基中sos3(?)降低更明显,而OE系和sOE萌发率无显著变化。
高盐土壤一般碱性较高,我们发现与WT相比,OE系在盐碱条件下的萌发率明显提高。
以上结果显示,TaCBL4和AtSOS3一样,也是钠、锂离子特异性响应基因,并且TaCBLA和AtSOS3在盐胁迫应答中发挥相似的功能,表明小麦中也存在SOS系统,而且不同植物间SOS系统存在保守性和通用性。本实验发现,TaCBL4在小麦抗钠离子育种中应该有针对性地设计在种子萌发阶段特异性表达,而在抗锉离子育种中则不受此限制。
有趣的是,低钾条件下,与Col-0相比,OE系和sOE系幼苗的地上部分明显变小,而AtSOS3过表达系则无显著变化,推测TaCBL4可能还调控钾离子通道的活性。这一结果表明,TaCBL4在盐胁迫应答中的作用与钾相关,幼苗阶段TaCBL4降低对Na+的耐性是由于负调控K+的转运引起的,显示了不同物种间SOS系统的特异性。
为了进一步认识TaCBL4的作用机制及其与AtSOS3功能的异同,我们利用酵母双杂交方法分离拟南芥中与TaCBL4相互作用的CIPK蛋白。结果发现,TaCBL4能与AtCIPK5/10/11/15/18/21/24这7个CIPKs互作,其中AtCIPK15/24为AtSOS3互作蛋白,而AtCIPK6/7不与TaCBL4互作。推测TaCBL4通过与AtCIPK11相互作用而抑制了后者活性,提高了植株的抗盐碱性,通过AtCIPK24(SOS2)互作而激活SOS通路促进Na+外排。TaCBL4和AtSOS3互作CIPKs种类存在相似性和特异性,而且TaCBL4和AtSOS3对与其共同互作的CIPKs的调控也可能存在差异性,那么这种特异性和差异性是否与TaCBL4过表达导致的低钾敏感型相关,需要进。步研究。
2、TaCBL3功能的初步分析
TaCBL3与水稻OsCBL6及拟南芥AtCBL2、AtCBL6相似性较高,其编码蛋白定位于液泡膜。离子和渗透胁迫下,TaCBL3过表达拟南芥株系与Col-0无明显差异。缺钾诱导TaCBL3的表达,而且缺钾条件下,与Col-0相比,TaCBL3过表达株系黄化更严重,植株生长抑制程度更显著。这表明,TaCBL3可能是钾离子通道的抑制因子,为植物离子转运机制研究提供了新的基因资源。
总之,本论文通过生物信息学和分子生物学方法,系统分析了小麦CBL和CIPK家族成员的序列和表达特征,通过CBL4的功能分析明确了不同物种间SOS通路对高盐、低钾响应存在通用性和特异性,为进一步认识SOS通路调控机制及离子胁迫响应特征奠定了基础,并为培育耐盐、钾营养高效作物新品种提供了分子元件。
Abiotic stress is one of major adverse constraints for crop yield. Modern molecular biology has been widely used for breeding, and a lot of stress tolerant crops were developed. In plants, the calcineurin B-like protein (CBL) family represents a unique group of calcium sensors and plays a key role in decoding calcium transients by specifically interacting with and regulating a family of protein kinases (CIPKs). Molecular genetics analyses of loss-of function mutants have implicated several CBL proteins and CIPKs as important components of abiotic stress responses. In Arabidopsis, the role of CBL and CIPK families in the response to abiotic stress has been largely elucidated; AtCBL4(SOS3) and AtCIPK24(SOS3) are two crucial component of SOS pathway, a ionic stress specific pathway in plants. However, the function of CBL and CIPK families as well as whether SOS pathway is present in wheat is still not clear, and whether SOS pathway is conserved in plants has not been documented.
Based on these scientific issues, we comprehensively analyzed the sequences of CBL and CIPK families, and isolated a set of TaCBL and TaCIPK genes from a wheat introgression cultivar SR3with high salt tolerance, and confirmed the role of two CBL genes TaCBL4and TaCBL3in the response to abiotic stress. The main research contents and results are summarized as follows:
1. The bioinfomatic analysis of wheat CBL and CIPK families and isolation of novel CBL/CIPK genes
According to the bioinformatic analysis, we electronically cloned14putative CBL genes and34putative CIPK genes from wheat. Multiple sequence alignment and phylogenetic tree analysis showed that these TaCBLs are much conserved in size and structure. Of them, five TaCBL proteins have N-myristoylation site and two possess transmembrane hydrophobic region, suggesting that these seven members play important roles in the stress response in wheat. TaCIPKs vary in sequence, but their second structure is very conserved. TaCIPKs from SR3were also difference from each other, which may possibly be resulted from gene duplication during evolution. Phylogenetic analysis showed that both TaCBLs and TaCIPKs had many paralogs, and they had higher similarity to homologues of rice than to those of Arabidopsis.
To know their salt-responsive patterns, these TaCIPKs were subject to BlastN against probes of our customized cDNA microarray. Among them, eight TaCIPKs had specifically matched probes, including four salt-inducible and four salt-restricted ones respectively; there had three salt-inducible probes, each of which referred to several TaCIPKs. These salt-responsive TaCIPKs may be involved in the response to salt stress, and their function is worthy of be elucidated.
According to these electronically cloned sequences, We cloned three TaCBLs and nine TaCIPKs for SR3. Based on their similarity to CBLs and CIPKs in Arabidopsis, these isolated genes were named as TaCBL1, TaCBL3, TaCBU, as well as TaCIPK2, TaCIPK8, TaCIPK15, TaCIPK17, TaCIPK19, TaCIPK30, TaCIPK32and TaCIPK34. Among TaCIPKs, TaCIPK8and TaCIPK15were up-regulated after exposure to salt, while TaCIPKI I was down-regulated.
2. The function analysis of wheat TaCBL4and TaCBL3
2.1TaCBL4structure and putative function analysis
TaCBL4possesses seven introns, and encodes a putative homolog of Arabidopsis CBL4/SOS3protein. TaCBL4localized in the cell membrane. TaCBL4transcribed in roots, leaves, stems, young panicle, awn, immature embryo and flag leaf, and it was drastically induced after exposure to200mM NaCl, suggesting its roles in development and response to salt stress.
To know its role in the response to abiotic stress, and its similarity to and difference from AtSOS3, TaCBL4was transformed into Arabidopsis Col-0wild type and sos3mutant to observe the phenotypic alteration, and AtSOS3overexpression Arabidopsis were used as the control. We found that TaCBL4ovevcxpression had no effect on vegetative and reproductive growth. In1/2MS medium, there had no difference among both roots and leaves of seedlings of Col-0(WT), TaCBL4overexpression (OE) lines, sos3overexpression TaCBL4(sOE)-Under NaCl treatment, in comparison with WT, sos3had shorter roots, and its leaves was severely bleached; both OE and sOE lines showed no significant difference, while there leaves became smaller, which phenocopied the effect of AtSOS3. This indicates that TaCBL4rescued the root phenotype of sos3, and firstly found that TaCBL4inhibited leaf growth under salt stress. Under LiCl treatment, in comparison with WT, sos3had shorter roots and smaller leaves, while OE and sOE lines as well as AtSOS3overexpression lines had longer roots and larger leaves, firstly demonstrating that TaCBL4and AtSOS3can enhance Li+tolerance. Under ABA and osmotic stress, no significant difference was found, indicating that TaCBL4is a salt specifically responsive gene.
In comparison with plant development, salt stress has a more severe adverse effect on seed germination. To know the role of TaCBL4at this stage, we compared the germination rate. Under NaCl treatment, sos3did not germinate, the germination rate of WT was reduced, while the germination rate of OE and sOE lines were comparable. Under LiCl treatment, the germination rate of WT and sos3was significantly lowered with a more extent in sos3, while that of OE and sOE lines was not affected; the growth of early-stage seedlings of WT and sos3was obviously inhibited, while the effect on OE and sOE lines was slight.
High saline soils are often alkaline. We found that under saline and alkaline conditions, OE lines had higher germination rate and superior growth ability at early stage than WT.
Above results indicate that, alike AtSOS3, TaCBL4was a Na+and Li+responsive gene, and they performed similar roles in the response to salt stress. This suggests that SOS pathway is under work in wheat, this pathway shares conservation among plants. Our work also shows that, it is a ideal strategy to let TaCBL4specifically transcribe at germination stage when breeding Na+tolerant wheat, and the gene can be constitutively expressed when breeding Li+tolerant cultivars.
Interestingly, under low K+conditions, in comparison with WT, the leaves of OE and sOE lines were smaller, while those of AtSOS3overexpression lines were similar, suggesting that TaCBL4can modulate the activities of K'channels. This result indicates that the role of TaCBL4in the response to salt was associated with K+ which outline the specificities of SOS pathway among different plants.
To further uncover the mechanism of TaCBL4performance and its similarity to and difference from AtSOS3, we isolated TaCBL4interacting AtCIPKs using the yeast two hybridization method. TaCBL4interacted with AtCIPK5,10,11,15,18,21and24, among which AtCIPK15and24are AtSOS3interacting proteins, while AICIPK6,7, the other AtSOS3interacting proteins, did not bind to TaCBL4. We speculate that TaCBL4interacts with AtCIPK11and inhibits its activity to improve the tolerance to saline and alkaline stress, and it interacts with AtCIPK24(SOS2) and triggers SOS pathway to accelerate Na+efflux. The coming question is whether or not the similarity and difference between TaCBL4and AtSOS3with respective to CIPK interaction, along with the putative difference between the modulation of TaCBL4and AtSOS3on their shared interacting CIPKs, are associated with the enhancement of TaCBL4-mediated sensitivity to low K+
2.2The primary analysis of TaCBL3function
TaCBL3shared high identities to rice OsCBL6and Arabidopsis AtCBL2and AtCBL6, and its encoding protein localized in vacuolar membrane. Under NaCl and osmotic stress, there had no phenotypic difference among Col-0and TaCBL3overexpression lines. Lack K'induced the expression of TaCBL3; under lack K+conditions, in comparison with Col-0, TaCBL3overexpression lines were more seriously etiolated, and their growth was more obviously inhibited. This demonstrated that TaCBL3is possibly a inhibitory factor of K'channels.
In summary, we comprehensively analyzed the sequence and expression characteristics of wheat CBL and CIPK families, and found the similarity and difference among SOS pathway of different plants during response to high Na+/Li+and low K+, which provided evidence for further understanding the regulatory mechanisms of SOS pathway and response to ionic stress, and these CBL and CIPK genes can be used for breeding new cultivars with salt tolerance and high K+utilization efficiency.
本研究基于以上科学问题,利用生物信息学方法系统分析了小麦CBL和CIPK基因家族,从本室培育的小麦渐渗系耐盐品种山融3号(SR3)中克隆了一系列TaCBLs和TaCIPKs基因,并初步验证了TaCBL4、TaCBL3在非生物胁迫应答中的功能。
一、小麦CBL和CIPK的生物信息学分析和新基因克隆
通过生物信息学分析,电子克隆了14个小麦CBL家族成员和34个小麦CIPK家族成员。多序列比对及进化树分析显示,这些TaCBLs在大小和结构上非常保守,其中5个具有豆蔻酰化位点,2个具有跨膜疏水区,暗示这7个TaCBLs在抗逆中发挥重要作用。TaCIPKs间序列差异比较大,但结构十分保守。SR3中TaCIPKs间也存在差异,这可能与基因倍增过程相关。进化树分析显示,小麦中TaCBLs和TaCIPKs都存在许多旁系同源基因,并且它们与水稻中同源基因的相似性高于拟南芥。
将这些电子克隆的TaCIPK基因与实验室定制的小麦cDNA芯片探针进行比对,以分析它们的盐胁迫应答特征。结果发现,8个TaCIPKs在芯片中有特异性匹配探针,其中4个为盐诱导表达基因,4个盐下调表达基因：另外,芯片中有3个探针分别对应多个TaCIPK序列,而且这3个探针在盐胁迫24h后表达量都显著提高。这些盐胁迫应答的TaCIPKs(?)可能都参与植物的盐胁迫应答反应,其功能需要进一步研究。
根据电子克隆序列,从SR3中克隆3个TaCBL和9个TaCIPK新基因,按照与拟南芥CBLs的同源性,分别命名为TaCBL1/3/4和TaC1PK2/8/11/15/17/19/30/32/34。 TaClPKs中,TaCIPK8、TaCIPK15在盐胁迫下上调表达,TaCIPK11下调表达,其他基因的应答模式有待进一步验证,为耐盐候选基因的鉴定及TaCBL互作蛋白的筛选奠定了基础。
二、小麦TaCBL4和TaCBL3功能分析
1、TaCBL4结构与可能作用分析
TaCBL4有7个内含子,其编码产物定位于细胞膜,与拟南芥同源基因AtSOS3(CBL4)相似,是AtSOS3同源基因。TaCBL4在小麦的根、叶、茎、幼穗、麦芒、幼胚、旗叶等组织中均能表达,NaCl(?)协迫下其表达量明显增加,表明TaCBL4在小麦生长发育和盐胁迫应答过程中均发挥重要作用。
为了验证TaCBL4(?)的生物学功能及其与AtSOS3(?)的异同,将其在拟南芥Col-0野生型和sos3突变体中异源表达,观察表型变化,并用AtSOS3在拟南芥中的过表达系作为对照。TaCBL4过表达不影响拟南芥的营养和生殖生长。在1/2MS培养基中,Col-0(WT)、TaCBL4过表达野生型系(OE)、TaCBL4过表达sos3(sOE)幼苗根长和地上部分无显著性差异。NaCl胁迫下,与WT相比,sos3幼苗的主根更短、地上部分漂白程度高,与朱建康实验室结果一致；而OE系和sOE系幼苗根与WT没有显著差别,但是叶片显著变小。这显示,TaCBL4能够恢复sos3根的表型,但首次发现OE系和sOE系在胁迫下抑制叶生长。LiCl胁迫下,与WT相比,sos3幼苗的主根明显变短、地上部分变小,而OH系、sOE系及AtSOS3过表达系幼苗正好相反,主根更长、地上部分更大,首次表明TaCBL4和AtSOS3过表达可以提高幼苗期锂离子胁迫抗性。在ABA处理和渗透胁迫下不同株系无明显差异,表明TaCBL4是一个离子特异性响应基因。
相比植株发育,盐胁迫对种子萌发的影响更大。为了分析TaCBL4在这阶段是否发挥重要作用,我们以绿色子叶张开率为标准,比较了不同株系在钠、锂离子胁迫下的萌发率差异。NaCl胁迫下,sos3不萌发,WT萌发率降低,而OE系和sOE萌发率受影响少,萌发率明显比WT高。LiCl处理下,WT和sos3的萌发率降低,基中sos3(?)降低更明显,而OE系和sOE萌发率无显著变化。
高盐土壤一般碱性较高,我们发现与WT相比,OE系在盐碱条件下的萌发率明显提高。
以上结果显示,TaCBL4和AtSOS3一样,也是钠、锂离子特异性响应基因,并且TaCBLA和AtSOS3在盐胁迫应答中发挥相似的功能,表明小麦中也存在SOS系统,而且不同植物间SOS系统存在保守性和通用性。本实验发现,TaCBL4在小麦抗钠离子育种中应该有针对性地设计在种子萌发阶段特异性表达,而在抗锉离子育种中则不受此限制。
有趣的是,低钾条件下,与Col-0相比,OE系和sOE系幼苗的地上部分明显变小,而AtSOS3过表达系则无显著变化,推测TaCBL4可能还调控钾离子通道的活性。这一结果表明,TaCBL4在盐胁迫应答中的作用与钾相关,幼苗阶段TaCBL4降低对Na+的耐性是由于负调控K+的转运引起的,显示了不同物种间SOS系统的特异性。
为了进一步认识TaCBL4的作用机制及其与AtSOS3功能的异同,我们利用酵母双杂交方法分离拟南芥中与TaCBL4相互作用的CIPK蛋白。结果发现,TaCBL4能与AtCIPK5/10/11/15/18/21/24这7个CIPKs互作,其中AtCIPK15/24为AtSOS3互作蛋白,而AtCIPK6/7不与TaCBL4互作。推测TaCBL4通过与AtCIPK11相互作用而抑制了后者活性,提高了植株的抗盐碱性,通过AtCIPK24(SOS2)互作而激活SOS通路促进Na+外排。TaCBL4和AtSOS3互作CIPKs种类存在相似性和特异性,而且TaCBL4和AtSOS3对与其共同互作的CIPKs的调控也可能存在差异性,那么这种特异性和差异性是否与TaCBL4过表达导致的低钾敏感型相关,需要进。步研究。
2、TaCBL3功能的初步分析
TaCBL3与水稻OsCBL6及拟南芥AtCBL2、AtCBL6相似性较高,其编码蛋白定位于液泡膜。离子和渗透胁迫下,TaCBL3过表达拟南芥株系与Col-0无明显差异。缺钾诱导TaCBL3的表达,而且缺钾条件下,与Col-0相比,TaCBL3过表达株系黄化更严重,植株生长抑制程度更显著。这表明,TaCBL3可能是钾离子通道的抑制因子,为植物离子转运机制研究提供了新的基因资源。
总之,本论文通过生物信息学和分子生物学方法,系统分析了小麦CBL和CIPK家族成员的序列和表达特征,通过CBL4的功能分析明确了不同物种间SOS通路对高盐、低钾响应存在通用性和特异性,为进一步认识SOS通路调控机制及离子胁迫响应特征奠定了基础,并为培育耐盐、钾营养高效作物新品种提供了分子元件。
Abiotic stress is one of major adverse constraints for crop yield. Modern molecular biology has been widely used for breeding, and a lot of stress tolerant crops were developed. In plants, the calcineurin B-like protein (CBL) family represents a unique group of calcium sensors and plays a key role in decoding calcium transients by specifically interacting with and regulating a family of protein kinases (CIPKs). Molecular genetics analyses of loss-of function mutants have implicated several CBL proteins and CIPKs as important components of abiotic stress responses. In Arabidopsis, the role of CBL and CIPK families in the response to abiotic stress has been largely elucidated; AtCBL4(SOS3) and AtCIPK24(SOS3) are two crucial component of SOS pathway, a ionic stress specific pathway in plants. However, the function of CBL and CIPK families as well as whether SOS pathway is present in wheat is still not clear, and whether SOS pathway is conserved in plants has not been documented.
Based on these scientific issues, we comprehensively analyzed the sequences of CBL and CIPK families, and isolated a set of TaCBL and TaCIPK genes from a wheat introgression cultivar SR3with high salt tolerance, and confirmed the role of two CBL genes TaCBL4and TaCBL3in the response to abiotic stress. The main research contents and results are summarized as follows:
1. The bioinfomatic analysis of wheat CBL and CIPK families and isolation of novel CBL/CIPK genes
According to the bioinformatic analysis, we electronically cloned14putative CBL genes and34putative CIPK genes from wheat. Multiple sequence alignment and phylogenetic tree analysis showed that these TaCBLs are much conserved in size and structure. Of them, five TaCBL proteins have N-myristoylation site and two possess transmembrane hydrophobic region, suggesting that these seven members play important roles in the stress response in wheat. TaCIPKs vary in sequence, but their second structure is very conserved. TaCIPKs from SR3were also difference from each other, which may possibly be resulted from gene duplication during evolution. Phylogenetic analysis showed that both TaCBLs and TaCIPKs had many paralogs, and they had higher similarity to homologues of rice than to those of Arabidopsis.
To know their salt-responsive patterns, these TaCIPKs were subject to BlastN against probes of our customized cDNA microarray. Among them, eight TaCIPKs had specifically matched probes, including four salt-inducible and four salt-restricted ones respectively; there had three salt-inducible probes, each of which referred to several TaCIPKs. These salt-responsive TaCIPKs may be involved in the response to salt stress, and their function is worthy of be elucidated.
According to these electronically cloned sequences, We cloned three TaCBLs and nine TaCIPKs for SR3. Based on their similarity to CBLs and CIPKs in Arabidopsis, these isolated genes were named as TaCBL1, TaCBL3, TaCBU, as well as TaCIPK2, TaCIPK8, TaCIPK15, TaCIPK17, TaCIPK19, TaCIPK30, TaCIPK32and TaCIPK34. Among TaCIPKs, TaCIPK8and TaCIPK15were up-regulated after exposure to salt, while TaCIPKI I was down-regulated.
2. The function analysis of wheat TaCBL4and TaCBL3
2.1TaCBL4structure and putative function analysis
TaCBL4possesses seven introns, and encodes a putative homolog of Arabidopsis CBL4/SOS3protein. TaCBL4localized in the cell membrane. TaCBL4transcribed in roots, leaves, stems, young panicle, awn, immature embryo and flag leaf, and it was drastically induced after exposure to200mM NaCl, suggesting its roles in development and response to salt stress.
To know its role in the response to abiotic stress, and its similarity to and difference from AtSOS3, TaCBL4was transformed into Arabidopsis Col-0wild type and sos3mutant to observe the phenotypic alteration, and AtSOS3overexpression Arabidopsis were used as the control. We found that TaCBL4ovevcxpression had no effect on vegetative and reproductive growth. In1/2MS medium, there had no difference among both roots and leaves of seedlings of Col-0(WT), TaCBL4overexpression (OE) lines, sos3overexpression TaCBL4(sOE)-Under NaCl treatment, in comparison with WT, sos3had shorter roots, and its leaves was severely bleached; both OE and sOE lines showed no significant difference, while there leaves became smaller, which phenocopied the effect of AtSOS3. This indicates that TaCBL4rescued the root phenotype of sos3, and firstly found that TaCBL4inhibited leaf growth under salt stress. Under LiCl treatment, in comparison with WT, sos3had shorter roots and smaller leaves, while OE and sOE lines as well as AtSOS3overexpression lines had longer roots and larger leaves, firstly demonstrating that TaCBL4and AtSOS3can enhance Li+tolerance. Under ABA and osmotic stress, no significant difference was found, indicating that TaCBL4is a salt specifically responsive gene.
In comparison with plant development, salt stress has a more severe adverse effect on seed germination. To know the role of TaCBL4at this stage, we compared the germination rate. Under NaCl treatment, sos3did not germinate, the germination rate of WT was reduced, while the germination rate of OE and sOE lines were comparable. Under LiCl treatment, the germination rate of WT and sos3was significantly lowered with a more extent in sos3, while that of OE and sOE lines was not affected; the growth of early-stage seedlings of WT and sos3was obviously inhibited, while the effect on OE and sOE lines was slight.
High saline soils are often alkaline. We found that under saline and alkaline conditions, OE lines had higher germination rate and superior growth ability at early stage than WT.
Above results indicate that, alike AtSOS3, TaCBL4was a Na+and Li+responsive gene, and they performed similar roles in the response to salt stress. This suggests that SOS pathway is under work in wheat, this pathway shares conservation among plants. Our work also shows that, it is a ideal strategy to let TaCBL4specifically transcribe at germination stage when breeding Na+tolerant wheat, and the gene can be constitutively expressed when breeding Li+tolerant cultivars.
Interestingly, under low K+conditions, in comparison with WT, the leaves of OE and sOE lines were smaller, while those of AtSOS3overexpression lines were similar, suggesting that TaCBL4can modulate the activities of K'channels. This result indicates that the role of TaCBL4in the response to salt was associated with K+ which outline the specificities of SOS pathway among different plants.
To further uncover the mechanism of TaCBL4performance and its similarity to and difference from AtSOS3, we isolated TaCBL4interacting AtCIPKs using the yeast two hybridization method. TaCBL4interacted with AtCIPK5,10,11,15,18,21and24, among which AtCIPK15and24are AtSOS3interacting proteins, while AICIPK6,7, the other AtSOS3interacting proteins, did not bind to TaCBL4. We speculate that TaCBL4interacts with AtCIPK11and inhibits its activity to improve the tolerance to saline and alkaline stress, and it interacts with AtCIPK24(SOS2) and triggers SOS pathway to accelerate Na+efflux. The coming question is whether or not the similarity and difference between TaCBL4and AtSOS3with respective to CIPK interaction, along with the putative difference between the modulation of TaCBL4and AtSOS3on their shared interacting CIPKs, are associated with the enhancement of TaCBL4-mediated sensitivity to low K+
2.2The primary analysis of TaCBL3function
TaCBL3shared high identities to rice OsCBL6and Arabidopsis AtCBL2and AtCBL6, and its encoding protein localized in vacuolar membrane. Under NaCl and osmotic stress, there had no phenotypic difference among Col-0and TaCBL3overexpression lines. Lack K'induced the expression of TaCBL3; under lack K+conditions, in comparison with Col-0, TaCBL3overexpression lines were more seriously etiolated, and their growth was more obviously inhibited. This demonstrated that TaCBL3is possibly a inhibitory factor of K'channels.
In summary, we comprehensively analyzed the sequence and expression characteristics of wheat CBL and CIPK families, and found the similarity and difference among SOS pathway of different plants during response to high Na+/Li+and low K+, which provided evidence for further understanding the regulatory mechanisms of SOS pathway and response to ionic stress, and these CBL and CIPK genes can be used for breeding new cultivars with salt tolerance and high K+utilization efficiency.
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