摘要
植物细胞分裂素激活的蛋白激酶(Mitogen-activated protein kinase,MAPK)是将外部信号转导到细胞内部的重要信号蛋白,参与植物的发育和多方面生理过程,包括病原物诱导的抗病反应。水稻MAPK基因功能的研究落后于模式植物拟南芥。Reyna和Yang(2006)根据水稻基因组序列推测基因组中存在17个MAPK成员,其中的9个MAPK基因参与水稻抗稻瘟病的抗病反应。
拟南芥的一个MAPK基因AtMPK4是系统获得性抗性(systemic acquired resistance,SAR)的负调控因子,介导细菌、真菌、病毒的广谱抗性。水稻中存在类似于拟南芥SAR的路径。本研究从明恢63中分离和鉴定了一个与AtMPK4基因同源的全长cDNA,基因命名为OsMPK6。由这个cDNA推导出的氨基酸序列与AtMPK4的同源性有84%,聚类分析显示其与AtMPK4和烟草中的一个MAPK基因NtMPK4在同一个分组中,暗示其可能与它们具相似的功能。表达谱分析显示OsMPK6在水稻的多个组织中组成型表达。
脱落酸(ABA)、赤霉素(GA3)和生长素(IAA)处理明恢63幼苗离体叶片,诱导OsMPK6表达量迅速上升,干旱胁迫也导致其mRNA积累,然而重金属盐(铜、镉和汞)以及高溶度的氯化钠和蔗糖并不影响其表达。低温(4℃)和高温(42℃)处理明恢63幼苗1小时和2小时,稍微诱导OsMPK6表达量上升。
抗病信号分子水杨酸(SA)、苯(1,2,3)噻二嗪-7-硫代羧酸-S-甲酯(BTH)、2,6-二氯异烟酸(INA)、茉莉酸(JA)和乙烯(ET),还有双氧水(H_2O_2)处理明恢63幼苗的离体叶片,5分钟就迅速诱导OsMPK6上升表达。用JA和SA共同处理水稻叶片诱导OsMPK6的表达,与它们各自单独处理并没有明显的区别。放线菌酮(CHX)也诱导OsMPK6上升表达,说明CHX是OsMPK6合成的负调节因子。
感病品种接种白叶枯菌与用水模拟接种相比,OsMPK6的表达量变化基本一致。抗病品种接种白叶枯菌,OsMPK6的表达量下降并大致维持比模拟接种更低的水平,携带不同R基因的抗病品种中表现出一样的趋势。明恢63接种不亲和稻瘟病小种V86013诱导OsMPK6上升表达。接种细菌和真菌病害诱导OsMPK6不同的表达量变化方式,暗示其可能在抗细菌和真菌病害中有着不同的调控方式。推测OsMPK6可能参与水稻的基本防卫反应,并受不同R基因的调节。
在水稻品种中花11中超量表达OsMPK6和采用RNA干扰技术(RNAi)抑制OsMPK6的表达都显著提高水稻对白叶枯病的抗性,抗病表型在T_1中与OsMPK6的表达量共分离。OsMPK6介导的抗性减少叶片中细菌的生长量,表明OsMPK6介导的抗病性是抗病方式而不是避病方式。OsMPK6介导的白叶枯病抗性广谱没有小种特异性,植株在苗期和成株期都表现为抗病。
超量表达OsMPK6提高水稻对白叶枯病的抗性显示OsMPK6正调控水稻抗病反应。超量表达植株抗病依赖于类病斑的形成。Northem分析表明植株出现类病斑时显著提高病程相关基因(pathogenesis-related genes)的表达量,组成型提高JA合成的重要基因OsAOS2表达,表明OsMPK6可能通过JA路径正调控抗病反应。植株没有类病斑时表现为感病,PR蛋白微弱表达。OsMPK6正调控的抗病性可能是局部抗性。
RNAi抑制OsMPK6/突变体也提高水稻对白叶枯病的抗性显示OsMPK6负调控水稻抗病反应。其沉默植株/突变体在成株期形成类病斑,接种后诱导抗病相关基因和PR基因的显著上升表达。SA合成途径的重要基因PAL诱导上升表达,OsMPK6可能通过SA路径负调抗病反应,调控的抗病性可能是系统抗性。OsMPK6沉默植株在不形成类病斑的苗期接种也表现为抗病,OsMPK6负调控的抗病性不依赖于类病斑。抑制OsMPK6的表达不影响创伤诱导JA合成的重要基因OsAOS2的上升表达。
虽然OsMPK6分别通过不同的途径正负调控水稻抗病反应,但是两种调控方式都通过WRKY03、NH1(like NPR1)、PR10和PR1a的途径,显示超量和抑制OsMPK6介导的抗病反应有一定的重叠。活性氧类物质(reactive oxygen species,ROS)参与了OsMPK6所调控的两条路径的抗病反应,这两条路径都导致水稻植株叶片上的细胞产生程序性死亡。超量和抑制OsMPK6的表达都提高SA在水稻植株中的含量,暗示OsMPK6可能是SA和JA途径的协调者。
在洋葱表皮细胞内瞬时表达的结果表明OsMPK6主要定位于细胞核中,在细胞质中也有微弱的表达。
Mitogen-activated protein kinase (MAPK) cascades play important roles in diversedevelopmental and physiological processes of plants, including pathogen-induced defenseresponses. On researching MAPKs in flee, they lag behind model plant Arabidopsis.Although at least 17 rice MAPKs have been identified and nine of these MAPK geneshave been shown to be pathogen or elicitor responsive, the exact role of most of theMAPKs in host-pathogen interaction is unknown.
AtMPK4 is an important negative gene of SAR in Arabidopsis, which confernon-specific resistance. Provious study showed there was a resistance pathway exists inrice as Arabidopsis. In this study, we isolated a rice homolog of AtMPK4 fromMinghui63, designed as OsMPK6. The deduced amino sequence from the full eDNAshared 84% sequence identity to AtMPK4 which showed the highest sequence similarity,Cluster analysis indicated it belong to the same group with AtMPK4 and NtMPK4. Theseresults hinted OsMPK6 would possess similar function as them. And OsMPK6constitutively expressed in multi tissues of rice.
OsMPK6 expression is induced rapidly by ABA, GA3, and IAA, drought stress alsoresulted in the accumulation of OsMPK6 transcripts. However, the heavy, metals, copper,cadmium and mercury, and high concentrations of NaCI and sucrose had no distinctinfluence on the expression of OsMPK6. Low(4℃) and high(42℃) temperature slightlyinduced OsMPK6 expression at 2 and 1 h after treatment, respectively.
As rapidly as 5 min after treatment, the plant defense signal molecules SA, BTH,INA, JA, ET and H_2O_2 induced the expression of OsMPK6. Comparing the expressionlevel of OsMPK6 after treatment with a single compound, interactions between SA, BTH,or INA and JA or ET showed no distinct effect on the expression of OsMPK6, CHX alsoinduced OsMPK6 expression at 60 to 120 min after treatment as compared to the control,which suggests that de novo synthesized negative regulator also may be involved in theregulation of OsMPK6 expression.
Comparing with the expression level of OsMPK6 in mock-inoculated plants,incompatible host-pathogen interaction further suppressed OsMPK6 expression andcompatible host-pathogen interaction slightly induced OsMPK6 expression. OsMPK6transcripts are induced and increased after inoculated blast. The difference expression ofOsMPK6 between bacterial blight and blast infection imply OsMPK6 could be functional in different way against bacterial and fungus pathogen and OsMPK6 would participate inbasic defense response, its expression was regulated with different R genes.
OsMPK6-overexpression and -suppression enhanced resistance to Xoo, theenhanced resistance cosegregated in T_1 family. OsMPK6-overexpression and-suppression plant conferred a broad spectrum of resistance to various Xoo strains, andshowed resistace to Xoo in rice full living stage.
OsMPK6 positively regulate resistance response. Overexpression of OsMPK6accompanying the development of lesion mimics on the leaves also enhanced theresistance to bacterial blight, whereas the OsMPK6-overexpressed plants that were free oflesion mimics were susceptible. Northern analysis results showed that differentialexpression of some defense-related genes in OsMPK6-overexpressed plants with orwithout lesion mimics, the resistant OsMPK6-overexpressed plants enhanced OsAOS2transcripts which is a important gene composed JA, explained OsMPK6 regulateresistance response by JA pathway, or local resistance.
OsMPK6 negatively regulate resistance response. OsMPK6-suppressedplants/mutant developed lesion mimics in adult stage, showed differential expression ofsome defense-related genes after inoculated Xoo. OsMPK6-suppressed plants enhancePAL expression that is key gene composed SA, explained OsMPK6 regulate resistanceresponse by SA pathway, or system resistance. OsMPK6-suppressed plants seedingwithout lesion mimics displayed resistance to Xoo, implied resistance does not require thedevelopment of lesion mimic. OsMPK6-suppressed plants don't change OsAOS2transcripts.
OsMPK6 both positively and negatively regulate resistance response using differentpathway. However, all of these resistant plants had elevated salicylic acid (SA) levels.And a subset of genes in the NH1 (an Arabidopsis NPR1 homologue) pathway showedenhanced expression, resulting in Program cell death and ROS. All the data suggestedOsMPK6 might be a coordinator between JA and SA pathway.
Observation of the onion epidermal strips under a confocal microscope showed thatOsMPK6 protein localized mainly in the nuclei and lightly in the cytoplasm
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