小金海棠转录因子MxFIT、MxIRO2的克隆及其与启动子关系的研究
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摘要
缺铁是限制果树特别是苹果树生长和果实品质的重要因素之一。多年来,人们围绕植物抗缺铁机理开展了许多研究,然而苹果抗缺铁的分子机理仍然不清楚。在实验室筛选到苹果铁高效基因型砧木小金海棠的基础上,本研究以小金海棠为试材,从中克隆到了两个受缺铁诱导表达的转录因子,命名为MxFTT和MxIRO2,分析及验证了其特性和功能,并研究了其与小金海棠铁吸收相关的两个重要功能基因MxIRT1和MxFRO2启动子之间的关系。研究结果为小金海棠铁吸收机制的深入挖掘奠定了基础。
根据苹果金冠基因组和已报道的与铁吸收相关转录因子的信息,克隆到了小金海棠MxFIT基因,其开放阅读框为966bp,编码321个氨基酸,属于螺旋-环-螺旋(bHLH)结构的转录因子,亚细胞定位分析表明MxFIT定位在细胞核,酵母单杂交表明MxFTT在酵母细胞中具有较强的激活能力。通过将MxFIT纯化蛋白注射新西兰大白兔获得了MxFIT抗体,利用实时荧光PCR和Western Blot分析了其在转录和翻译水平的表达,MxFIT主要在小金海棠根系中表达,在叶片中几乎不表达。在根中,缺铁不仅可以强烈诱导MxFIT基因表达,而且可以诱导MxFIT蛋白积累,说明MxFTT在转录和翻译水平上均受缺铁调控。在拟南芥中过表达MxFIT可使转基因植株在缺铁条件下增强铁的吸收应答反应,提高了植株的抗缺铁能力。因此,MxFIT在缺铁应答反应中起着重要的作用。
同时克隆到了另一个bHLH家族转录因子MxIRO2,该转录因子基因的开放阅读框为762bp,编码253个氨基酸。亚细胞定位显示MxIRO2定位在细胞核,MxIRO2和GAL4DNA结合域的融合蛋白在酵母中能激活LacZ报告基因的表达,其激活能力较弱,可被1mM3-AT抑制。通过将纯化的MxIRO2蛋白注射新西兰大白兔获得了MxIR02的抗体,利用实时荧光PCR和Western Blot分析了其在转录和翻译水平的表达,结果表明MxIRO2在小金海棠的根部和叶部均受缺铁诱导表达,在转录和翻译水平的表达均先升高后降低。
酵母双杂交表明MxFIT和MxIRO2之间存在互作关系,利用烟草注射瞬时表达方法发现MxFIT和MxIR02同时注射烟草时可激活MxIRT1和MxFRO2启动子；将含启动子删除片段的农杆菌注射烟草表明MxIRTl启动子在-485～-236bp处存在与MxFIT和MxIRO2转录因子结合的序列,MxFRO2启动子在-813～-564bp处存在与MxFIT和MxIR02转录因子结合的序列。通过转化原生质体方法检测了MxIRT1和MxFRO2启动子的活性, MxIRT1启动子在-764～-485bp处存在增强启动子活性的元件,MxFRO2启动子在-1704～-1228bp和-564--329bp处存在增强启动子活性的元件。
本研究从转录因子与功能基因启动子的关系入手证明了MxFIT和MxIRO2互作后可调控MxIRTl和MxFRO2,同时利用转基因拟南芥也证明了IRT1和FRO2是FIT调控的下游功能基因。因此,MxFTT和MxIRO2是小金海棠铁吸收调控过程中的重要转录因子。
Iron deficiency is one of the important factors that affect the yield and quality in fruit trees, especially for apple tree. People have conducted a lot of researches about the mechanism of resistance to iron deficiency for years. However, the molecular mechanism of resistance to iron deficiency in apple tree is still not very clear. Malus xiaojinensis (Cheng et Jiang) is the first apple genotype identified as being resistant to iron deficiency by our research team. In this study, two iron-deficiency-induced transcription factors, MxFIT and MxIRO2, were cloned in M. xiaojinensis and the characteristics and functions were also analysed. MxIRTl and MxFRO2had been proved as two major genes related to iron uptake. The relationship of the promoters of MxIRTl and MxFR02with MxFIT and MxIR02transcription factors was studied in this paper. This work offers an important basis for further investigating the mechanism of fruit tree adaptation to iron-deficiency stress.
According to the genome of Golden Delicious and the transcriptome sequencing of M. xiaojinensis, an iron deficiency response transcription factor gene, MxFIT, was cloned from M. xiaojinensis. MxFIT encoded a basic helix-loop-helix protein and contained a966bp open reading frame. MxFIT protein was targeted to the nucleus in onion epidermal cells and showed strong transcriptional activation in yeast cells. The antibody was obtained by injecting a New Zealand rabbit with the purified MxPIT-His fusion protein. Real-time PCR and Western blot analysis revealed that MxFIT was up-regulated in roots under iron deficiency at both mRNA and protein levels, while almost no expression was detected in leaves irrespective of iron supply. Ectopic expression of MxFIT resulted in enhanced iron deficiency responses in Arabidopsis under iron deficiency and stronger resistance to iron deficiency. Thus, MxFIT might be involved in iron uptake and plays an important role in iron deficiency response.
In this study, another bHLH iron-related transcription factor IRO2, containing a762bp open reading frame and encoding253amino acids, had been cloned from M. xiaojinensis. The MxIRO2protein was targeted to nucleus in onion epidermal cells and had weaker transcription activation ability in yeast cells. The activation degree was restrained by1mM3-AT. The antibody was obtained by injecting a New Zealand rabbit with the purified MxIR02-His fusion protein. The spatiotemporal expression of MxIR02under iron deficiency was then assayed. The results indicated that MxIR02was induced in both roots and leaves when subject to iron deficiency. The expression model in transcription and translation level was increased at first and then decreased with iron deficiency stress.
Yeast two-hybrid analysis proved that MxFIT could interact with MxIRO2. The promoters of MxIRTl and MxFRO2were activated in the tobacco leaves by coinfiltrated with agrobacterium containing MxFIT and MxIRO2. The binding sequences with MxFIT and MxIRO2transcription factor might locate-485～-236bp in MxIRT1promoter and-813～-564bp in MxFR02promoter. There might be reinforcing elements in-764～-485bp of MxIRT1promoter and-1704～-1228bp and-564～-329bp of MxFOR2promoter.
Therefore, based on the relationship between transcription factors and functional gene promoters, we proved that MxFIT and MxIRO2could regulate MxIRTl and MxFRO2after interactions. Meanwhile, we found IRT1and FRO2were regulated by FIT in transgenic Arabidopsis. MxFTT and MxDRO2were two important transcription factors in regulation of iron absorption in Malus xiaojinensis.
根据苹果金冠基因组和已报道的与铁吸收相关转录因子的信息,克隆到了小金海棠MxFIT基因,其开放阅读框为966bp,编码321个氨基酸,属于螺旋-环-螺旋(bHLH)结构的转录因子,亚细胞定位分析表明MxFIT定位在细胞核,酵母单杂交表明MxFTT在酵母细胞中具有较强的激活能力。通过将MxFIT纯化蛋白注射新西兰大白兔获得了MxFIT抗体,利用实时荧光PCR和Western Blot分析了其在转录和翻译水平的表达,MxFIT主要在小金海棠根系中表达,在叶片中几乎不表达。在根中,缺铁不仅可以强烈诱导MxFIT基因表达,而且可以诱导MxFIT蛋白积累,说明MxFTT在转录和翻译水平上均受缺铁调控。在拟南芥中过表达MxFIT可使转基因植株在缺铁条件下增强铁的吸收应答反应,提高了植株的抗缺铁能力。因此,MxFIT在缺铁应答反应中起着重要的作用。
同时克隆到了另一个bHLH家族转录因子MxIRO2,该转录因子基因的开放阅读框为762bp,编码253个氨基酸。亚细胞定位显示MxIRO2定位在细胞核,MxIRO2和GAL4DNA结合域的融合蛋白在酵母中能激活LacZ报告基因的表达,其激活能力较弱,可被1mM3-AT抑制。通过将纯化的MxIRO2蛋白注射新西兰大白兔获得了MxIR02的抗体,利用实时荧光PCR和Western Blot分析了其在转录和翻译水平的表达,结果表明MxIRO2在小金海棠的根部和叶部均受缺铁诱导表达,在转录和翻译水平的表达均先升高后降低。
酵母双杂交表明MxFIT和MxIRO2之间存在互作关系,利用烟草注射瞬时表达方法发现MxFIT和MxIR02同时注射烟草时可激活MxIRT1和MxFRO2启动子；将含启动子删除片段的农杆菌注射烟草表明MxIRTl启动子在-485～-236bp处存在与MxFIT和MxIRO2转录因子结合的序列,MxFRO2启动子在-813～-564bp处存在与MxFIT和MxIR02转录因子结合的序列。通过转化原生质体方法检测了MxIRT1和MxFRO2启动子的活性, MxIRT1启动子在-764～-485bp处存在增强启动子活性的元件,MxFRO2启动子在-1704～-1228bp和-564--329bp处存在增强启动子活性的元件。
本研究从转录因子与功能基因启动子的关系入手证明了MxFIT和MxIRO2互作后可调控MxIRTl和MxFRO2,同时利用转基因拟南芥也证明了IRT1和FRO2是FIT调控的下游功能基因。因此,MxFTT和MxIRO2是小金海棠铁吸收调控过程中的重要转录因子。
Iron deficiency is one of the important factors that affect the yield and quality in fruit trees, especially for apple tree. People have conducted a lot of researches about the mechanism of resistance to iron deficiency for years. However, the molecular mechanism of resistance to iron deficiency in apple tree is still not very clear. Malus xiaojinensis (Cheng et Jiang) is the first apple genotype identified as being resistant to iron deficiency by our research team. In this study, two iron-deficiency-induced transcription factors, MxFIT and MxIRO2, were cloned in M. xiaojinensis and the characteristics and functions were also analysed. MxIRTl and MxFRO2had been proved as two major genes related to iron uptake. The relationship of the promoters of MxIRTl and MxFR02with MxFIT and MxIR02transcription factors was studied in this paper. This work offers an important basis for further investigating the mechanism of fruit tree adaptation to iron-deficiency stress.
According to the genome of Golden Delicious and the transcriptome sequencing of M. xiaojinensis, an iron deficiency response transcription factor gene, MxFIT, was cloned from M. xiaojinensis. MxFIT encoded a basic helix-loop-helix protein and contained a966bp open reading frame. MxFIT protein was targeted to the nucleus in onion epidermal cells and showed strong transcriptional activation in yeast cells. The antibody was obtained by injecting a New Zealand rabbit with the purified MxPIT-His fusion protein. Real-time PCR and Western blot analysis revealed that MxFIT was up-regulated in roots under iron deficiency at both mRNA and protein levels, while almost no expression was detected in leaves irrespective of iron supply. Ectopic expression of MxFIT resulted in enhanced iron deficiency responses in Arabidopsis under iron deficiency and stronger resistance to iron deficiency. Thus, MxFIT might be involved in iron uptake and plays an important role in iron deficiency response.
In this study, another bHLH iron-related transcription factor IRO2, containing a762bp open reading frame and encoding253amino acids, had been cloned from M. xiaojinensis. The MxIRO2protein was targeted to nucleus in onion epidermal cells and had weaker transcription activation ability in yeast cells. The activation degree was restrained by1mM3-AT. The antibody was obtained by injecting a New Zealand rabbit with the purified MxIR02-His fusion protein. The spatiotemporal expression of MxIR02under iron deficiency was then assayed. The results indicated that MxIR02was induced in both roots and leaves when subject to iron deficiency. The expression model in transcription and translation level was increased at first and then decreased with iron deficiency stress.
Yeast two-hybrid analysis proved that MxFIT could interact with MxIRO2. The promoters of MxIRTl and MxFRO2were activated in the tobacco leaves by coinfiltrated with agrobacterium containing MxFIT and MxIRO2. The binding sequences with MxFIT and MxIRO2transcription factor might locate-485～-236bp in MxIRT1promoter and-813～-564bp in MxFR02promoter. There might be reinforcing elements in-764～-485bp of MxIRT1promoter and-1704～-1228bp and-564～-329bp of MxFOR2promoter.
Therefore, based on the relationship between transcription factors and functional gene promoters, we proved that MxFIT and MxIRO2could regulate MxIRTl and MxFRO2after interactions. Meanwhile, we found IRT1and FRO2were regulated by FIT in transgenic Arabidopsis. MxFTT and MxDRO2were two important transcription factors in regulation of iron absorption in Malus xiaojinensis.
引文
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Li L, Cheng X, Ling HQ. Isolation and characterization of Fe (III)-chelate reductase gene LeFRO1 in tomato. Plant Molecular Biology,2004,54:125-136
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Landsberg EC. Regulation of iron-stress-response by whole-plant activity. Journal of Plant Nutrition, 1984,7:609-621
Landsberg, EC. Function of rhizodermal transfer cells in Fe stress response mechanism of Capsicum annuum L. Plant Physiology,1986,82:511-517
Li CJ, Zhu XP, Zhang FS. Role of shoot in regulation of iron deficiency responses in cucumber and bean plants. Journal of Plant Nutrition,2000,23:1809-1818
Li L, Cheng X, Ling HQ. Isolation and characterization of Fe (III)-chelate reductase gene LeFRO1 in tomato. Plant Molecular Biology,2004,54:125-136
Li XX, Duan XP, Jiang HX, et al. Genome-wide analysis of basic/helix-loop-helix transcription factor family in Rice and Arabidopsis, Plant Physiology,2006,141:1167-1184
Ling HQ, Bauer P, Bereczky Z, et al. The tomato fer gene encoding a bHLH protein controls iron-uptake responses in roots. Proceedings of the National Academy of Sciences of the United States of America,2002,99:13938-13943
Lingam P, Mohrbacher J, Brumbarova T, et al. Interaction between the bHLH transcription factor FIT and ETHYLENE INSENSITIVE3/ETHYLENE INSENSITIVE3-LIKE1 reveals molecular linkage between the regulation of iron acquisition and ethylene signaling in Arabidopsis. Plant Cell,2011, 23:1815-29
Long TA, Tsukagoshi H, Busch W, et al. The bHLH transcription factor POPEYE regulates response to iron deficiency in Arabidopsis roots. Plant Cell,2010,22:2219-2236
Maas FM, van de Wetering DAM, van Beusichem ML, et al. Characterization of phloem iron and its possible role in the regulation of Fe-efficiency reaction. Plant Physiology,1988,87:167-171
Massari ME, Murre C. Helix-loop-helix proteins:regulators of transcription in Eucaryotic organisms. Molecular and Cellular Biology,2000,20:429-440
Meiser J, Lingam S, Bauer P. Posttranslational regulation of the iron deficiency basic helix-loop-helix transcription factor FIT is affected by iron and nitric oxide. Plant Physiology,2011,157: 2154-2166
Mermod N, O' Neill EA, Kelly TJ, et al. The proline-rich transcriptional activator of CTF-NF1 is distinct from the replication and DNA binding domain. Cell,1989,58:741-753
Michelet B, Lukaszewicz M, Dupriez V, et al. A plant plasma membrane proton-ATPase gene is regulated by development and environment and shows signs of translational regulation. Plant Cell, 1994,6:1375-1389
Morsomme P, Boutry M. The plant plasma membrane H+-ATPase:Structure, function and regulation. Biochimica et Biophysica Acta-Biomembranes,2000,1465:1-16
Murgia I, Tarantino D, Soave C, et al. Arabidopsis CYP82C4 expression is dependent on Fe availability and circadian rhythm, and correlates with genes involved in the early Fe deficiency response. Journal of Plant Physiology,2011,168:894-902
Murre C, McCaw PS, Vaessin H, et al. Interactions between heterologous helix-loop-helix proteins generate complexes that bind specifically to a common DNA sequence. Cell,1989,58(3):537-544
Niebur WS, Fehr WR. Agronomic evaluation of soybean genotypes resistant to iron deficiency chlorosis. Crop Science,1981,21:551-554
Niu X, Guiltinan MJ. DNA binding specificity of the wheat bZIP protein EmBP-1. Nucleic Acids Research,1994,22:4969-4978
Ogo Y, Kobayashi T, Itai RN, et al. A novel NAC transcription factor IDEF2 that recognizes the iron deficiency-responsive element 2 regulates the genes involved in iron homeostasis in plants. Journal of Biological Chemistry,2008,283:13407-13417
Ogo Y, Itai RN, Nakanishi H, et al. Isolation and characterization of IRO2, a novel iron-regulated bHLH transcription factor in graminaceous plants. Journal of Experimental Botany,2006, 57:2867-2878
Ogo Y, Itai RN, Kobayashi T, et al. OsIRO2 is responsible for iron utilization in rice and improves growth and yield in calcareous soil. Plant Molecular Biology,2011,75:593-605
Ogo Y, Itai RN, Nakanishi H, et al. The ricebHLHprotein OsIRO2 is an essential regulator of the genes involved in Fe uptake under Fe-deficient conditions. Plant Jounery,2007,51:366-377
Palmgren MG. Plant plasma membrane H+-ATPases:powerhouses for nutrient uptake. Annual Review of Plant Physiology and Plant Molecular Biology.2001,52:817-845.
Qi JN, Yu SC, Zhang FL, et al. Reference gene seletion for real-time quantitative polymerase chain reaction of mRNA transcript levels in Chinese cabbage (Brassica rapa L. ssp. pekinensis). Plant Molecular Biology Reporter,2010,28:597-604
Robinson NJ Groom SJ, Groom QJ. The froh gene family from Arabidopsis thaliana:putative iron-chelate reductase. Plant and Soil,1997,196:245-248
Robinson NJ, Procter CM, Connolly EL, et al. A ferric-chelate reductase for iron uptake from soils. Natrue,1999,397:649-697
Romera FJ, Alcantara E, Guardia de la MD. Ethylene production by Fe-deficient roots and its involvement in the regulation of Fe-deficiency stress responses by strategy I plants. Annals of Botany,1999,83:51-55
Romera FJ, Welch RM, Norvell WA, et al. Iron requirement for and effects of promoters and inhibitors of ethylene action or stimulation of Fe (III)-chelate reductase in roots of Strategy I species. Biometals,1996,9(1):45-50
Romheld V, Marschner H. Mobilization of iron in the rhizosphere of different plant species. Journal of Plant Nutrition,1986,2:155-192
Santi S, Schmidt W. Dissecting iron deficiency-induced proton extrusion in Arabidopsis roots. New Phytologist,2009,183:1072-1084
Schmidt W. Iron solutions:acquisition strategies and signaling pathway in plants. Trends in Plant Science,2003,8:188-193
Sivitz A, Grinvalds C, Barberon M, et al. Proteasome-mediated turnover of the transcriptional activator FIT is required for plant iron-deficiency responses. Plant Journal,2011,66:1044-1152
Staiger D. Chemical strategies for iron acquisition in plants. Angewandte Chemie International Edition, 2002,41(13):2259-2264
Struhl K. Helix-turn-helix, zinc-finger, and leucine-zipper motifs for eukaryotic transcriptional regulatory proteins. Trends in Biochemical Sciences,1989,14:137-140
Takahashi M, Yamaguchi H, Nakanishi H, et al. Cloning two genes for nicotianamine aminotransferase, a critical enzyme in iron acquisition (Strategy II) in graminaceous plants. Plant Physiology,1999, 121:947-956
Tang WH, Zhang JL, Wang ZY, et al. The cause of deviation made in determining the molecular weight of His-tag fusion proteins by SDS-PAGE. Acta Phytophysiologica Sinica,2000,26:64-68
Tjaden G, Coruzzi GM. A novel AT-rich DNA binding protein that combines an HMG-like DNA binding domain with a putative transcription domain. Plant Cell,1994,6:107-111
Velasco R, Zharkikh A, Affourtit J, et al. The genome of the domesticated apple (Malus×domestica Borkh.). Nature genetics,2010,42:833-839
Vert G, Grotz N, Dedald6champ F,et al. IRT1, an Arabidopsis transporter essential for iron uptake from the soil and for plant growth. Plant Cell,2002,14:1223-1233
Vorwieger A, Gryczka C, Czihal A, et al. Iron assimilation and transcription factor controlled synthesis of riboflavin in plants. Planta,2007,226:147-158
Walker EL, Connolly EL. Time to pump iron:iron-deficiency-signaling mechanisms of higher plants. Current Opinion in Plant Biology,2008,11:1-6
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