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拟南芥β-葡萄糖苷酶调节ABA含量动态变化与耐旱性关系研究
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摘要
植物激素ABA不仅参与调控种子休眠和萌发过程,还广泛参与干旱、盐碱和冷害等多种非生物胁迫应答反应。这些生理功能的实现依赖于细胞内ABA水平动态且快速的精细调控。细胞内ABA含量受复杂的从头合成途径、羟基化分解代谢和共价结合失活的共同控制。ABA葡萄糖酯(ABA-GE)是植物体内ABA结合失活的主要形式,这种结合态的ABA-GE能否重新释放出具生物学活性的游离ABA,目前所知甚少。
     我们筛选了功能获得性ABA不敏感突变体gai6,该突变基因编码β-葡萄糖苷酶BGLU10。bglu10的T-DNA插入纯合突变体气孔无论胁迫与否均趋于较大的开放状态,导致水分丧失过快而不耐干旱;相反BGLU10的超表达株系比野生型耐旱性更强,而且ABA、NaCl及甘露醇等胁迫处理时BGLU10的超表达株系比野生型萌发后子叶变绿数量更少,而bglu10突变体则表现更好的萌发后幼苗生长态势。无论正常浇水还是干旱处理条件下BGLU10的超表达株系中ABA含量均高于野生型WT,而bglu10突变体中ABA含量却比野生型稍低。ABA或干旱反应的标志性基因的表达分析表明,脱水处理可诱导RAB18、RD29B和RD22的表达显著上调,而且BGLU10-OE植物中此三个基因的表达量高于WT,而bglu10突变体中该三个基因的表达量则低于WT; ABA响应的转录因子ATHB6同样表现出类似的表达模式。此外,干旱处理条件下植物体内总β-葡萄糖苷酶活性与植物的耐旱性呈正相关。这些结果均暗示BGLU10可能通过调控ABA含量进而使干旱耐性基因表达量升高而增强植物的耐早性。
     带GFP标签的BGLU10融合蛋白及RFP与液泡膜标志蛋白γ-TIP1融合蛋白的原生质体瞬时表达的荧光共定位结果显示,BGLU10酶蛋白定位于液泡中;且我们的实验也显示拟南芥总β-葡萄糖苷酶活性的最适pH值为6.0左右,与已知的液泡酸碱环境相吻合。BGLU10基因组织表达的GUS染色结果表明该基因在植物各组织包括根、茎、叶、花及果荚均有表达,尤其是成熟的输导组织中表达更为强烈。此外ABA、盐害、脱水及冷胁迫均可诱导BGLU10基因的表达上调,暗示了β-葡萄糖苷酶BGLU10可能参与了多种胁迫下游离ABA的水解释放与植物的胁迫耐性反应。
     总之,本研究使人们对植物体内ABA水平的动态调节机制有了新的认识,即ABA的从头合成、羟基化分解代谢、共价结合失活及结合态ABA的重新酶解释放,协同精细调控植物体内游离ABA含量,使植物体得以应对复杂多变的胁迫环境和生长发育需要。
The phytohormone ABA plays crucial roles in various physiological processes including seed dormancy, germination and adaptive responses to environmental stress such as drought, cold and high salt in soil. The content fluctuate of cellular ABA levels allow plants to adapt to the changing physiological and environmental conditions. Cellular ABA is controlled by complex de novo synthesis pathways, hydroxylation and conjugation reactions. ABA glucose ester (ABA-GE) is the dominating form covalent bounding ABA in plants. However, the issue of whether the inactive ABA-GE could re-release and produce the biological active ABA remains to be clarified.
     We screened a gain-function of ABA-insensitive mutant gai6, the mutation gene encodedβ-glucosidase BGLU10. T-DNA insertion homozygous mutant bglulO showed faster withering and death than WT under drought stress, and stomatal behavior analysis showed that stomata of bglu10 tend to keeping bigger aperure than that of WT, which leads to greater loss of water whether under stress or not. By contrast, the over-expression lines of BGLU10 showed stronger drought-tolerance than WT. In addition, BGLU10-OE lines behaved fewer green cotyledons after germination than WT, while the mutant bglu10 performed better situation after germination and seedling growth under ABA, NaCl and mannitol stress treatments. Determination of ABA content in plants under both normal watering and drought condition showed BGLU10-OE lines hold higher ABA content than that in WT, but ABA content in bglu10 slightly lower than that in WT. The results of expression analyses for marker genes responding to ABA and/or drought showed RAB18, RD29B and RD22 were dramatically induced by dehydration stress, and it worthy noted that the expressions level of these genes were highest in BGLU10-OE lines but lowest in bglulO mutant. The similar expression pattern was found for homeodomain-leucine zipper transcription factor gene ATHB6, which was known significantly up-regulated by ABA. Furthermore, the total P-glucosidase activity is positive correlate to the drought resistance in bglu10 mutant, WT and BGLU10-OE plants. These results also effectively explained the difference between ABA content and drought-tolerance of different plants, suggesting BGLU 10 may have the function of hydrolysing ABA-GE and releasing of free ABA to enhance stress-tolerance response of plants.
     Fluorescence co-localization result of BGLU10::GFP and y-TIP 1::RFP (a tonoplast marker protein) fusion proteins using transient expression in protoplast of mesophyll cells showed BGLU 10 enzyme protein localized in vacuole, and our experiment also showed the optimal pH is about pH 6.0 of totalβ-glucosidase activity in Arabidopsis, which matched the known vacuolar pH environment. GUS staining analysis of BGLU10 gene organ-specific expression showed that this gene is transcripted in various organizations, including roots, stems, leaves, flowers and siliqua, especially in mature conducting tissues. In addition, BGLU10 was induced by ABA, salt, drought and cold stress, suggesting thatβ-glucosidase BGLU 10 may be involved in variety of stresses and hydrolysis of ABA-GE producing free ABA in plant stress tolerance response.
     In conclusion, this study provide us a new understanding of ABA level dynamic adjustment mechanism in plants, namely, by fine controlling of free ABA content through collaboratively tuning pathways of ABA de novo synthesis, hydroxylation catabolism, covalent conjugation and releasing free ABA from ABA-GE, so plants can cope with complex and changing stress environment and the developing needs.
引文
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