2个玉米光敏色素C基因的转录丰度对多种光质处理的响应
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摘要
【目的】通过NCBI数据库获得2个玉米PHYC及相关数据,并进行相关生物信息学分析。利用实时荧光定量PCR(q RT-PCR)分析2个玉米PHYC在玉米各器官的转录丰度,以及其转录丰度对多种光质处理、黑暗到各种光质转换和光周期处理(长日照和短日照)的响应,为研究玉米PHYC在玉米幼苗去黄化与开花期的调控机制奠定基础。【方法】采用玉米B73自交系为研究材料,通过RT-PCR分别对Zm PHYC1和Zm PHYC2的全长ORF序列进行克隆;借助相关软件对其进行生物信息学分析,利用q RT-PCR分析这两个基因在玉米各器官中的转录丰度,及其转录丰度对各种光照处理的响应。【结果】Zm PHYC1和Zm PHYC2的全长ORF均为3 408 bp,编码1 135个氨基酸基序,分子量分别为126.14和126.07 k D。生物信息学分析表明,玉米phy C蛋白可以分为6个功能区段:节奏周期蛋白—Ah核转运接受蛋白—专一蛋白区段(Per-Arnt-Sim,PAS)、c GMP受激磷酸二酯酶区段(GAF)、色素区段(PHY)和PAS相关区段(PRD,包含2个PAS区段)、组氨酸激酶A区段和组氨酸激酶ATP酶区段,但是Zmphy C2在PRD区段仅有一个PAS区段。氨基酸水平的系统发育树分析表明,Zmphy C1和Zmphy C2与禾本科物种phy C有很高的一致性,且与甘蔗和高粱phy C的亲缘关系较近。q RT-PCR分析表明,Zm PHYC1和Zm PHYC2的表达在根和叶中的转录丰度均较高,同时对持续蓝光和白光响应强烈;在黑暗到各种光质转换处理中,这两个PHYC的表达模式相似。在黑暗转到远红光、红光、蓝光和白光的0.5 h,Zm PHYC1和Zm PHYC2的转录表达均急剧上升,随后迅速下降到自身起始黑暗时的水平以下,并上下波动。这两个基因对长日照和短日照的光周期处理也能积极响应,在长日照条件下,2个Zm PHYC出现了极其相似的表达模式,均在光照和黑暗阶段各出现1个峰值;在短日照条件下,这两个基因的表达模式差异较大,Zm PHYC1的峰值出现在进入黑暗后6 h,而Zm PHYC2的峰值出现在进入光照阶段2 h。【结论】玉米phy C蛋白可以分成6个功能区段,但是Zmphy C2在PRD区段仅具有一个PAS相关区段。2个玉米PHYC转录丰度具有组织特异性。在各种光质处理中,Zm PHYC1和Zm PHYC2的表达模式相近,可能二者存在功能冗余,在转录水平上前者的丰度高于后者,推测Zm PHYC1在玉米中起更重要的作用,并且可能二者在功能上存在分工。Zm PHYC1和Zm PHYC2对各种光质和光周期处理均有较强的响应,推测二者在调控玉米光形态建成和开花中具有重要作用。
【Objective】To study the functions of phytochrome C genes in seedling de-etiolation and flowering regulation in maize(Zea mays L.), two phytochrome C genes of maize(Zm PHYC1 and Zm PHYC2) were selected from the NCBI database and analyzed by bioinformatic methods. The transcription abundances of two Zm PHYC genes was analyzed in different tissues and in response to light qualities, transitions from the dark to different light conditions, photoperiod treatment(long day and short day) by quantitative RT-PCR(q RT-PCR).【Method】B73 inbred line was used in this study, the full length ORFs of two Zm PHYC genes were cloned by RT-PCR. The proper clones were sequenced and analyzed by bioinformatics software. The transcription abundances of two Zm PHYC genes in different tissues and in response to light treatments was detected using q RT-PCR.【Result】Both the full length ORFs of Zm PHYC1 and Zm PHYC2 contained 3408 nucleotides and encoded two polypeptides with 1135 amino acid motifs, and their molecular weight was 126.14 k D and 126.07 k D, respectively. Two Zmphy C proteins were able to be further divided into six domains: Per(period circadian protein)-Arnt(Ah receptor nuclear translocator protein)-Sim(single-minded protein)(PAS), c GMP-stimulated phosphodiesterase(GAF), phytochrome(PHY), PAS-related domain(PRD) containing two PAS, His Kinase A domain(His KA), Histidine kinase-like ATPases(HATPase_c), while Zmphy C2 lacked a PAS in PRD domain. Phylogenetic analysis indicated that the two Zmphy C proteins belonged to the same branch with phy C proteins from other graminaceous species, especially with the phy C proteins from sugarcane and sorghum. q RT-PCR assays showed that both Zm PHYC1 and Zm PHYC2 belonged to tissue-specific genes and highly expressed in roots and leaves. The transcription abundances of the both genes were very high under blue and white light conditions. Both Zm PHYC1 and Zm PHYC2 displayed similar expression patterns during transitions from the dark to different light conditions. The transcription abundances of the both genes went dramatically up at 0.5 h after transitions from the dark to far-red, red, blue, or white light. And then they quickly dropped and waved below their own levels in the dark. Both Zm PHYC1 and Zm PHYC2 were also able to respond to long-day or short-day treatments. During long-day treatment, they likely had one peak in either light or dark period. During short-day treatment, they showed different expression patterns, the peak of Zm PHYC1 happened at 6 h after conversion to dark period, while Zm PHYC2 occurred at 2 h after conversion to light period.【Conclusion】 Zmphy C1 protein kept six domains, while Zmphy C2 lacks a PAS in PRD domain. The transcription abundances of the both PHYC genes were tissue-specific in maize. Similar expression patterns of Zm PHYC1 and Zm PHYC2 genes in response to different light treatments suggest that they both might keep redundant functions. Since the transcription abundances of Zmphy C1 were higher than these of Zmphy C2, Zmphy C1 might have more important role in seedling responding to light than Zmphy C2, which may be due to the existence of different functions in maize. Both Zm PHYC1 and Zm PHYC2 respond to light and photoperiod treatments, suggesting that they are involved in seedling de-etiolation and flowering time control in maize. Thus their roles in crop improvement are worthy of more exploration in the future.
【Objective】To study the functions of phytochrome C genes in seedling de-etiolation and flowering regulation in maize(Zea mays L.), two phytochrome C genes of maize(Zm PHYC1 and Zm PHYC2) were selected from the NCBI database and analyzed by bioinformatic methods. The transcription abundances of two Zm PHYC genes was analyzed in different tissues and in response to light qualities, transitions from the dark to different light conditions, photoperiod treatment(long day and short day) by quantitative RT-PCR(q RT-PCR).【Method】B73 inbred line was used in this study, the full length ORFs of two Zm PHYC genes were cloned by RT-PCR. The proper clones were sequenced and analyzed by bioinformatics software. The transcription abundances of two Zm PHYC genes in different tissues and in response to light treatments was detected using q RT-PCR.【Result】Both the full length ORFs of Zm PHYC1 and Zm PHYC2 contained 3408 nucleotides and encoded two polypeptides with 1135 amino acid motifs, and their molecular weight was 126.14 k D and 126.07 k D, respectively. Two Zmphy C proteins were able to be further divided into six domains: Per(period circadian protein)-Arnt(Ah receptor nuclear translocator protein)-Sim(single-minded protein)(PAS), c GMP-stimulated phosphodiesterase(GAF), phytochrome(PHY), PAS-related domain(PRD) containing two PAS, His Kinase A domain(His KA), Histidine kinase-like ATPases(HATPase_c), while Zmphy C2 lacked a PAS in PRD domain. Phylogenetic analysis indicated that the two Zmphy C proteins belonged to the same branch with phy C proteins from other graminaceous species, especially with the phy C proteins from sugarcane and sorghum. q RT-PCR assays showed that both Zm PHYC1 and Zm PHYC2 belonged to tissue-specific genes and highly expressed in roots and leaves. The transcription abundances of the both genes were very high under blue and white light conditions. Both Zm PHYC1 and Zm PHYC2 displayed similar expression patterns during transitions from the dark to different light conditions. The transcription abundances of the both genes went dramatically up at 0.5 h after transitions from the dark to far-red, red, blue, or white light. And then they quickly dropped and waved below their own levels in the dark. Both Zm PHYC1 and Zm PHYC2 were also able to respond to long-day or short-day treatments. During long-day treatment, they likely had one peak in either light or dark period. During short-day treatment, they showed different expression patterns, the peak of Zm PHYC1 happened at 6 h after conversion to dark period, while Zm PHYC2 occurred at 2 h after conversion to light period.【Conclusion】 Zmphy C1 protein kept six domains, while Zmphy C2 lacks a PAS in PRD domain. The transcription abundances of the both PHYC genes were tissue-specific in maize. Similar expression patterns of Zm PHYC1 and Zm PHYC2 genes in response to different light treatments suggest that they both might keep redundant functions. Since the transcription abundances of Zmphy C1 were higher than these of Zmphy C2, Zmphy C1 might have more important role in seedling responding to light than Zmphy C2, which may be due to the existence of different functions in maize. Both Zm PHYC1 and Zm PHYC2 respond to light and photoperiod treatments, suggesting that they are involved in seedling de-etiolation and flowering time control in maize. Thus their roles in crop improvement are worthy of more exploration in the future.
引文
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[8]MONTE E,ALONSO J M,ECKER J R,ZHANG Y L,LI X,YOUNG J,PHILLIPS S A,QUAIL P H.Isolation and characterization of phy C mutants in Arabidopsis reveals complex crosstalk between phytochrome signaling pathways.The Plant Cell,2003,15(9):1962-1980.
[9]FANKHAUSER C,CASAL J J.Phenotypic characterization of a photomorphogenic mutant.The Plant Journal,2004,39(5):747-760.
[10]BATSCHAUER A.Photoreceptors of higher plants.Planta,1998,206(4):479-492.
[11]LI J G,LI G,WANG H Y,DENG X W.Phytochrome signaling mechanisms.The Arabidopsis Book,2011,9:e0148.
[12]BAE G,CHOI G.Decoding of light signals by plant phytochromes and their interacting proteins.Annual Review of Plant Biology,2008,59:281-311.
[13]SAKAMOTO K,NAGATANI A.Nuclear localization activity of phytochrome B.The Plant Journal,1996,10(5):859-868.
[14]廖祥儒,史海水,尚丹,韩国辉.植物中的光敏色素.生物技术通讯,2004,15(1):95-97.LIAO X R,SHI H S,SHANG D,HAN G H.Photochromes in plants.Letters in Biotechnology,2004,15(1):95-97.(in Chinese)
[15]REED J W,NAGATANI A,ELICH T D,FAGAN M,CHORY J.Phytochrome A and phytochrome B have overlapping but distinct functions in Arabidopsis development.Plant Physiology,1994,104(4):1139-1149.
[16]SHARROCK R A,CLACK T.Heterodimerization of type II phytochromes in Arabidopsis.Proceedings of the National Academy of Sciences of the USA,2004,101(31):11500-11505.
[17]SHARROCK R A,CLACK T.Patterns of expression and normalized levels of the five Arabidopsis phytochromes.Plant Physiology,2002,130(1):442-456.
[18]CLACK T,MATHEWS S,SHARROCK R A.The phytochrome apoprotein family in Arabidopsis,is encoded by five genes:The sequences and expression of PHYD,and PHYE.Plant Molecular Biology,1994,25(3):413-427.
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