光敏色素B调控的一个水稻NBS-LRR基因的表达和功能初探
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摘要
利用水稻基因芯片比较野生型、phyA和phyB突变体受连续红光照射后的基因表达图谱,从中筛选到一个受phyB介导的红光信号特异诱导的、编码NBS-LRR类蛋白的基因,命名为PB-LRR (phyB-regulated NBS-LRR)。
为了揭示PB-LRR基因在水稻生长发育中作用,本研究首先对该基因表达的器官特异性、对不同激素的应答反应进行了分析。结果表明,PB-LRR基因在叶中表达水平最高,在茎中最低。光对PB-LRR基因表达的诱导是由phyB调控的,与phyA无关。此外PB-LRR基因的表达受外源ABA的抑制,并且受环境因子的调控；不受PEG处理的影响,但很可能与光周期有关。本研究将构建好的PB-LRR启动区：GUS植物表达载体并转化至野生型和phyB突变体中,GUS染色结果表明,在phyB突变体转基因植株中的GUS活性明显低于野生型,而且GUS活性主要出现在叶片上部。这些结果表明PB-LRR基因的表达不仅受phyB介导的光信号调控,而且受发育调控。
为了进一步了解PB-LRR基因的功能,本研究通过农杆菌愈伤组织侵染法成功培育出了该基因的过量表达和缺失表达的转基因株系。对生长14d的两种转基因株系的幼苗进行表型分析发现过量表达的株系主根较野生型和缺失表达的株系长,但地上部分各株系的差异不明显。由此推测该基因的过量表达可以使主根更为发达。另外,干旱处理后,PB-LRR基因的缺失表达株系的耐旱性比野生型和过量表达株系明显要差,野生型和过量表达株系耐旱性相当。这一结果说明,该基因与水稻的干旱胁迫耐性相关。
生物信息学分析发现,PB-LRR基因属于CC-NBS-LRR类基因,目前报道的这类基因多数为抗性基因,因而推测其与水稻的抗病性相关。本研究中利用Western blotting对茉莉酸(JA)和水杨酸(SA)处理后的野生型和过量表达的株系进行了PR-1蛋白表达水平的分析,并未发现明显差异,说明PB-LRR基因可能不参与JA、SA依赖的PR-1蛋白参与的抗病信号途径。
本研究为进一步分析PB-LRR基因在phyB介导的光反应和水稻生长发育中的作用奠定了基础。
We compared gene profiles among wild-type, phyA and phyB mutants grown under continuous red light or in the dark. One gene, which encodes NBS-LRR family protein, is specifically up-regulated by red light perceived by phyB. We named this gene PB-LRR (phyB-regulated NBS-LRR).
To access the biological functions of PB-LRR gene in rice growth, we analyzed the organ-specificity of PB-LRR expression as well as effects of exogenous hormones on PB-LRR expression. The level of PB-LRR transcripts was highest in leaf and lowest in shoot. Our results showed that light-induced expression of PB-LRR gene was attributed to phyB, not phyA. Moreover, expression of PB-LRR gene was inhibited by exogenous ABA treatment, and also regulated by environmental factors. The level of PB-LRR transcripts was not affected by PEG but probably influenced by photoperiod. The expression cassette of putative promoter region of PB-LRR and GUS (ProPB-LRR:GUS) was constructed and introduced into wild-type (WT) and phyB mutant, respectively. GUS activities were obviously detected in upper part of leaves in the WT background, but only weakly detected in the phyB mutant background. These results demonstrate that PB-LRR gene is not only regulated by phyB-mediated light signals but also by leaf development in rice.
In order to analyze the PB-LRR role in rice in detail, we produced two kinds of transgenic lines:PB-LRR OE/WT (overexpression) and PB-LRR RNAi/WT (RNA interference) by Agroinfection method. Comparison of phenotypes of 14-day-old wild-type and PB-LRR transgenic seedlings revealed that the seminal roots of PB-LRR OE/WT were significantly longer than wild-type and PB-LRR RNAi/WT. No obvious difference was observed in the above-ground parts of wild type and PB-LRR transgenic lines. In addition, PB-LRR RNAi/WT showed obvious increased sensitivity to drought stress. The drought tolerance of PB-LRR OE/WT lines was similar to wild-type plants. These results suggest that PB-LRR gene is involved in regulating the rice drought tolerance.
Bioinformatics analysis revealed that PB-LRR gene belonged to CC-NBS-LRR gene family. In this family, most members have been reported to be resistance gene. Thus, it was deduced that PB-LRR gene probably play a role in rice disease resitance. However, the PR-1 protein levels have no difference in wild-type and PB-LRR OE/WT leaves treated with exogenous JA and SA using western blotting analysis. These results suggest that the deduced disease resistance of PB-LRR has no relationship with expression of PR-1 proteins induced by either JA- or SA- defense pathway.
Our results underlie the knowledge for further understanding the roles of PB-LRR in phyB-mediated signaling, disease resistance and drought tolerance in rice.
为了揭示PB-LRR基因在水稻生长发育中作用,本研究首先对该基因表达的器官特异性、对不同激素的应答反应进行了分析。结果表明,PB-LRR基因在叶中表达水平最高,在茎中最低。光对PB-LRR基因表达的诱导是由phyB调控的,与phyA无关。此外PB-LRR基因的表达受外源ABA的抑制,并且受环境因子的调控；不受PEG处理的影响,但很可能与光周期有关。本研究将构建好的PB-LRR启动区：GUS植物表达载体并转化至野生型和phyB突变体中,GUS染色结果表明,在phyB突变体转基因植株中的GUS活性明显低于野生型,而且GUS活性主要出现在叶片上部。这些结果表明PB-LRR基因的表达不仅受phyB介导的光信号调控,而且受发育调控。
为了进一步了解PB-LRR基因的功能,本研究通过农杆菌愈伤组织侵染法成功培育出了该基因的过量表达和缺失表达的转基因株系。对生长14d的两种转基因株系的幼苗进行表型分析发现过量表达的株系主根较野生型和缺失表达的株系长,但地上部分各株系的差异不明显。由此推测该基因的过量表达可以使主根更为发达。另外,干旱处理后,PB-LRR基因的缺失表达株系的耐旱性比野生型和过量表达株系明显要差,野生型和过量表达株系耐旱性相当。这一结果说明,该基因与水稻的干旱胁迫耐性相关。
生物信息学分析发现,PB-LRR基因属于CC-NBS-LRR类基因,目前报道的这类基因多数为抗性基因,因而推测其与水稻的抗病性相关。本研究中利用Western blotting对茉莉酸(JA)和水杨酸(SA)处理后的野生型和过量表达的株系进行了PR-1蛋白表达水平的分析,并未发现明显差异,说明PB-LRR基因可能不参与JA、SA依赖的PR-1蛋白参与的抗病信号途径。
本研究为进一步分析PB-LRR基因在phyB介导的光反应和水稻生长发育中的作用奠定了基础。
We compared gene profiles among wild-type, phyA and phyB mutants grown under continuous red light or in the dark. One gene, which encodes NBS-LRR family protein, is specifically up-regulated by red light perceived by phyB. We named this gene PB-LRR (phyB-regulated NBS-LRR).
To access the biological functions of PB-LRR gene in rice growth, we analyzed the organ-specificity of PB-LRR expression as well as effects of exogenous hormones on PB-LRR expression. The level of PB-LRR transcripts was highest in leaf and lowest in shoot. Our results showed that light-induced expression of PB-LRR gene was attributed to phyB, not phyA. Moreover, expression of PB-LRR gene was inhibited by exogenous ABA treatment, and also regulated by environmental factors. The level of PB-LRR transcripts was not affected by PEG but probably influenced by photoperiod. The expression cassette of putative promoter region of PB-LRR and GUS (ProPB-LRR:GUS) was constructed and introduced into wild-type (WT) and phyB mutant, respectively. GUS activities were obviously detected in upper part of leaves in the WT background, but only weakly detected in the phyB mutant background. These results demonstrate that PB-LRR gene is not only regulated by phyB-mediated light signals but also by leaf development in rice.
In order to analyze the PB-LRR role in rice in detail, we produced two kinds of transgenic lines:PB-LRR OE/WT (overexpression) and PB-LRR RNAi/WT (RNA interference) by Agroinfection method. Comparison of phenotypes of 14-day-old wild-type and PB-LRR transgenic seedlings revealed that the seminal roots of PB-LRR OE/WT were significantly longer than wild-type and PB-LRR RNAi/WT. No obvious difference was observed in the above-ground parts of wild type and PB-LRR transgenic lines. In addition, PB-LRR RNAi/WT showed obvious increased sensitivity to drought stress. The drought tolerance of PB-LRR OE/WT lines was similar to wild-type plants. These results suggest that PB-LRR gene is involved in regulating the rice drought tolerance.
Bioinformatics analysis revealed that PB-LRR gene belonged to CC-NBS-LRR gene family. In this family, most members have been reported to be resistance gene. Thus, it was deduced that PB-LRR gene probably play a role in rice disease resitance. However, the PR-1 protein levels have no difference in wild-type and PB-LRR OE/WT leaves treated with exogenous JA and SA using western blotting analysis. These results suggest that the deduced disease resistance of PB-LRR has no relationship with expression of PR-1 proteins induced by either JA- or SA- defense pathway.
Our results underlie the knowledge for further understanding the roles of PB-LRR in phyB-mediated signaling, disease resistance and drought tolerance in rice.
引文
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[2]Nagy F, Schafer E. Phytochromes control photomorphogenesis by differentially regulated, interacting signaling pathways in higher plants [J]. Annu Rev Plant Biol,2002,53:329-355.
[3]Casal JJ. Phytochromes, cryptochromes, phototropin:photoreceptor interactions in plants [J]. Photochem Photobiol,2000,71:1-11.
[4]Briggs WR, Christie JM. Phototropins 1 and 2:versatile plant blue-light receptors [J]. Trends Plant Sci,2002,7:204-210.
[5]Ulm R, Nagy F. Signaling and gene regulation in response to ultraviolet light [J]. Curr Opin Plant Biol,2005,8:477-482.
[6]Quail PH. Photosensory perception and signaling in plant cells:new paradigms [J]? Curr Opin Cell Biol,2002,14:180-188.
[7]Nagatani A, Reed JW, Chory J. Isolation and initial characterization of Arabidopsis mutants that are deficient in phytochrome A [J]. Plant Physiol,1993,102:269-277.
[8]Parks BM, Quail PH. hy8, a new class of Arabidopsis long hypocotyl mutants deficient in functional phytochrome A [J]. Plant Cell,1993,5:39-48.
[9]Reed JW, Nagpal P, Poole DS, et al. Mutations in the gene for the red-far-red light receptor phytochrome B alter cell elongation and physiological responses throughout Arabidopsis development[J]. Plant Cell,1993,5:147-157.
[10]Devlin PF, Patel SR, Whitelam GC. Phytochrome E influences internode elongation and flowering time in Arabidopsis [J].Plant Cell,1998,10:1479-1487.
[11]Devlin PF, Robson PR, Patel SR, et al. Phytochrome D acts in the shade-avoidance syndrome in Arabidopsis by controlling elongation growth and flowering time [J]. Plant Physiol,1999,119: 909-915.
[12]刘圈炜,何云,齐胜利,等.光敏色素研究进展[J].中国农学通报,2005,2：237-241.
[13]Quail P H. An emerging map of the phytochromes [J]. Plant Cell Environ,1997,20:657-665.
[14]Matsushita T, Mochizuki N, Nagatani A. Dimers of the N-terminal domain of phytochrome B are functional in the nucleus [J].Nature,2003,424:571-574.
[15]Bae G, Choi G. Decoding of light signals by plant phytochromes and their interacting proteins [J]. Annu. Rev. Plant Biol,2008,59:281-311.
[16]Lincoln T, Eduarclo Z. Plant Physiology [M].Sunderland:Sinauer Associates,2002,Inc. pp. 376-376.
[17]Andel F, Lagarias JC, Mathies RA. Resonance Raman analysis of chromophore structure in the Lumi-R photoproduct of phytochrome [J]. Biochem,1996,35:15997-16008.
[18]Gu X, Chen Z, Zhu Y. Phytochrome and photoregulation [J]. Avta Bot Sin,1997,39:675-681.
[19]Kendrick RE, Kronenberg G. Photomorphogenesis in Plants [M],2nd edn. Dordreeht:Kluwer,1994, pp.707-707.
[20]顾建伟,刘婧,薛彦久,等.光敏色素在水稻生长发育中的作用[J].中国水稻科学,2011,25：130-135.
[22]王静,王艇.高等植物光敏色素的分子结构、生理功能和进化特征[J].植物学通报,2007,24：649-658.
[23]Deng XW, Matsici M, Wei N, et al. COP1, an Arabidopsis regulatory gene, encodes a protein with both a zinc-binding motify and a G beta homologous domain [J]. Cell,1992,71:791-801.
[24]Van VE. Leaf expansion—an integration plant behaviour [J]. Plant J,1998,3:583-590.
[25]Keara AF, Garry CW (1999). Phytochromes and shade-avoidance responses in plants [J]. Plant Cell Envion,1999,22:1463-1473.
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