玉米低磷应答转录因子ZmPHR1的克隆及功能分析
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摘要
玉米(Zea mays L.)是世界上重要的粮食、经济和饲料兼用型作物。大部分玉米种植于热带和温带的酸性或碱性土壤中,而磷素作为植物重要的营养元素之一极易被酸性土壤中的Fe3+、Al3+以及碱性土壤中的Ca2+结合形成难溶性化合物,导致土壤中有效磷缺乏,最终严重影响玉米的产量和品质。尽管磷肥的施用能够缓解土壤中有效磷缺乏的问题,但是随之而来的是引起更多磷素在土壤中被固定、流失、水体富营养化以及磷矿资源的缺乏等一系列问题。研究表明,只有极少数玉米自交系属磷高效利用型。所以,培育低磷条件下的磷高效利用型玉米种质为育种所用,是我们迫切需要解决的问题。磷的利用效率是一个受多基因控制的数量性状,导入单个功能基因并不能达到改善某一个数量性状的效果,而单个转录因子的导入可以调控下游多个基因的表达,因此,将转录因子作为研究对象以期提高玉米的磷利用效率。到目前为止,磷利用相关转录因子的研究以拟南芥和水稻中AtPHR1和OsPHR2研究最为详尽,二者都属于MYB-CC转录因子家族,而且在进化上同源,且已被证明是磷饥饿响应的重要调控因子,在植物磷积累调控中起重要作用。然而,在玉米中,尚未有与PHR1类似的转录因子的报道。
从筛选到的磷高效利用型玉米材料中克隆与AtPHR1同源的基因,并对该基因的序列特征、特性以及功能进行了分析研究。主要结论如下：
1.对36份玉米材料进行低磷胁迫处理,采用影响磷利用效率的两个关键因素干物质量和磷积累量的绝对值和相对值进行综合评价,并辅助以生理生化特性分析,最终筛选出磷高效利用型玉米材料478和太系113。
2.以AtPHR1为基础序列,利用序列相似性分析方法,从磷高效利用型玉米材料478中获得一个氨基酸序列。生物信息学分析表明,该序列编码一个含有449个氨基酸的蛋白,与AtPHR1、OsPHR1和OsPHR2的一致性分别为43%、64%和45%:具有与AtPHR1.OsPHR1和OsPHR2一样高度保守的MYB-DNA结合域和CC结构域,属于一个新MYB-CC转录因子家族,命名为ZmPHR1。系统进化树分析表明,2mPHR1与AtPHRl,OsPHR1,OsPHR2同属一个亚类,相比AtPHR1,与OsPHR1、 OsPHR2的亲缘关系更近一些。ZmPHR1基因位于玉米第1条染色体长臂上。基因结构分析表明,ZmPHR1具有7个外显子和6个内含子,与AtPHR1和OsPHR1的基因结构相似。
3.利用基因枪法将含有ZmPHR1::GFP融合蛋白的植物表达载体导入洋葱表皮细胞。结果表明,仅在核内观察到GFP荧光,说明ZmPHR1在核内表达,为一个MYB-CC转录因子。RT-PCR分析表明,无论高磷还是低磷培养,ZmPHRl在不同组织器官中均有表达,在叶中表达量最高,根中次之,茎中最弱,属组织特异性表达。qPC(?)进一步分析表明,随着磷饥饿处理时间的延长,ZmPHR1的表达量处在一个动态变化中,存在一个正负反馈调节。
4.通过农杆菌浸染花蕾法将含有ZmPHR1::GFP的植物双元表达载体导入拟南芥,通过潮霉素抗性筛选、DNA-PCR、RT-PCR和GFP检测,获得过量表达的转基因拟南芥株系。
5.通过对拟南芥ZmPHR1过表达株系和野生型株系的表型对比分析得出,相比野生型株系,ZmPHR1的过量表达明显改善转基因植株的生长。特别是在低磷条件下,相比野生型株系,过表达株系的茎叶干重和无机磷含量均显著提高。
6.通过对拟南芥磷饥饿响应基因的表达分析得出,在低磷条件下,相比野生型株系,ZmPHR1过表达株系的根和茎叶中的AtIPS1、AtPS2、AtPS3和AtRNS1的表达均上调,并且同时上调了根中磷转运蛋白基因Pht1;2、Pht1;8和Pht1;9的表达,推测这些基因的上调是植株生物量和无机磷含量提高的原因。
Maize (Zea mays L.) is an important food, economic and forage crop in the world. It is grown widely in both acidic and calcareous soils, where inorganic phosphate (Pi) is easily formed insoluble complexes with aluminum and iron in acidic conditions or calcium ions in alkaline soil. Therefore, maize yield and quality is limited by the poor availability of Pi. Although the application of phosphorus (P) fertilizer could alleviate the problem of lack of Pi, a series of attendant problems are caused including phosphorus fixation, erosion in soils, eutrophication of water body and phosphate rock resources shortage. Many researches have showed that there are only a few of maize inbreds with high phosphorus use efficiency (PUE). Producing more maize germplasm with high PUE is an urgent problem to be resolved. PUE is a complex multi-gene trait, with a single gene contributing only a small percentage to the phenotype. A single transcription factor (TF) frequently coordinates the expression of multiple genes in response to environmental signals and regulate polygene quantitative traits. So, TFs would be used to regulate maize PUE. So far, two TFs related to Pi starvation were studied most detailed, which are AtPHRl in Arabidopsis and OsPHR2in rice. They belong to the MYB-CC TF family and are evolutionary homology. Both AtPHRl and OsPHR2were verified to be important regulators for phosphate starvation response and play important roles in the regulation of plant phosphorus accumulation. However, little is known about the functions of PHR-like TFs in maize.
In this study, a homolog gene sequence of AtPHRl was isolated from an inbred line of maize with high PUE. The charicaterization and functions the gene were studied. The results are summarized as follows:
1. Thirty-six maize materials were treated by low phosphorus stress. The absolute values and relative values of dry matter amount and phosphorus accumulate amount were used to be an evaluating indicator to screen high PUE genotypes, which are assisted with analysis of physiological-biochemical charicteristic. Two high PUE genotypes478and TX113were obtained.
2. According to the sequence alignment analysis, a homolog gene sequence of AtPHR1was isolated from an inbred line478of maize with high PUE. Bioinformatics analysis showed that the sequence encodes a449-amino acid protein. Multiple sequence alignment results showed that the predicted amino acids sequence shared43%,64%, and45%identity with the amino acid sequences of AtPHR1, OsPHRl and OsPHR2, respectively. The results also indicated that the four amino acid sequences were highly conserved in both MYB DNA-binding domain (BD) and CC domain. So, the sequence designated as ZmPHRl. Phylogenetic analysis showed that ZmPHRl was clustered in the same group as AtPHRl, OsPHR1and OsPHR2. ZmPHRl was sub-grouped with OsPHR1and OsPHR2, which was consistent with their close evolutionary relationship. ZmPHR1was located on long arm of maize chromosome1. The gene structure of ZmPHR1had a similar splicing pattern as AtPHRl, OsPHRl with seven exons and six introns.
3. The construct with ZmPHR1::GFP fusion protein was introduced into onion epidermal cells by particle bombardment. The fluorescence signal was located predominately in the nuclei. The results demonstrated that ZmPHR1is a nuclear protein and is a MYB-CC TF.ZmPHR1showed a tissue specificity expression pattern, with the highest expression in leaves, the higher expression in roots and the weakest expression in stems. Further qPCR analysis showed that, ZmPHRl expression of different tissues was in a dynamic change along with the prolonging of phosphate starvation treatment time, and exist a positive and negative feedback to phosphate starvation.
4. A binary vector containing the ZmPHRl::GFP fusion protein was constructed and introduced into Arabidopsis by floral dip method. The transgenic Arabidopsis lines with overexpressing ZmPHR1were obtained and verified by Hygromycin resistant screening, DNA-PCR, RT-PCR and GFP.
5. The phenotype of ZmPHRl overexpression lines and wild type (WT) Arabidopsis analysis showed that overexpression of ZmPHRl could significantly improve the growth of transgenic plants. The dry matter yield and Pi content of ZmPHR1-overexpressing lines were much higher than those of WT, especially under LP conditions.
6. The expression patterns of Pi-starvation-inducible genes were examined in ZmPHR1-overexpressing Arabidopsis plants. Under LP conditions, compared to wild-type plants, expression of AtIPS1, AtPS2, AtPS3and AtRNSl in shoots and roots was up-regulated in ZmPHR1overexpression lines, and Pht1;2、Phtl;8and Phtl;9were also up-regulated in roots. It is inferred that the upregulation of these Pi-starvation-inducible genes could increase the plant biomass and phosphorus content.
从筛选到的磷高效利用型玉米材料中克隆与AtPHR1同源的基因,并对该基因的序列特征、特性以及功能进行了分析研究。主要结论如下：
1.对36份玉米材料进行低磷胁迫处理,采用影响磷利用效率的两个关键因素干物质量和磷积累量的绝对值和相对值进行综合评价,并辅助以生理生化特性分析,最终筛选出磷高效利用型玉米材料478和太系113。
2.以AtPHR1为基础序列,利用序列相似性分析方法,从磷高效利用型玉米材料478中获得一个氨基酸序列。生物信息学分析表明,该序列编码一个含有449个氨基酸的蛋白,与AtPHR1、OsPHR1和OsPHR2的一致性分别为43%、64%和45%:具有与AtPHR1.OsPHR1和OsPHR2一样高度保守的MYB-DNA结合域和CC结构域,属于一个新MYB-CC转录因子家族,命名为ZmPHR1。系统进化树分析表明,2mPHR1与AtPHRl,OsPHR1,OsPHR2同属一个亚类,相比AtPHR1,与OsPHR1、 OsPHR2的亲缘关系更近一些。ZmPHR1基因位于玉米第1条染色体长臂上。基因结构分析表明,ZmPHR1具有7个外显子和6个内含子,与AtPHR1和OsPHR1的基因结构相似。
3.利用基因枪法将含有ZmPHR1::GFP融合蛋白的植物表达载体导入洋葱表皮细胞。结果表明,仅在核内观察到GFP荧光,说明ZmPHR1在核内表达,为一个MYB-CC转录因子。RT-PCR分析表明,无论高磷还是低磷培养,ZmPHRl在不同组织器官中均有表达,在叶中表达量最高,根中次之,茎中最弱,属组织特异性表达。qPC(?)进一步分析表明,随着磷饥饿处理时间的延长,ZmPHR1的表达量处在一个动态变化中,存在一个正负反馈调节。
4.通过农杆菌浸染花蕾法将含有ZmPHR1::GFP的植物双元表达载体导入拟南芥,通过潮霉素抗性筛选、DNA-PCR、RT-PCR和GFP检测,获得过量表达的转基因拟南芥株系。
5.通过对拟南芥ZmPHR1过表达株系和野生型株系的表型对比分析得出,相比野生型株系,ZmPHR1的过量表达明显改善转基因植株的生长。特别是在低磷条件下,相比野生型株系,过表达株系的茎叶干重和无机磷含量均显著提高。
6.通过对拟南芥磷饥饿响应基因的表达分析得出,在低磷条件下,相比野生型株系,ZmPHR1过表达株系的根和茎叶中的AtIPS1、AtPS2、AtPS3和AtRNS1的表达均上调,并且同时上调了根中磷转运蛋白基因Pht1;2、Pht1;8和Pht1;9的表达,推测这些基因的上调是植株生物量和无机磷含量提高的原因。
Maize (Zea mays L.) is an important food, economic and forage crop in the world. It is grown widely in both acidic and calcareous soils, where inorganic phosphate (Pi) is easily formed insoluble complexes with aluminum and iron in acidic conditions or calcium ions in alkaline soil. Therefore, maize yield and quality is limited by the poor availability of Pi. Although the application of phosphorus (P) fertilizer could alleviate the problem of lack of Pi, a series of attendant problems are caused including phosphorus fixation, erosion in soils, eutrophication of water body and phosphate rock resources shortage. Many researches have showed that there are only a few of maize inbreds with high phosphorus use efficiency (PUE). Producing more maize germplasm with high PUE is an urgent problem to be resolved. PUE is a complex multi-gene trait, with a single gene contributing only a small percentage to the phenotype. A single transcription factor (TF) frequently coordinates the expression of multiple genes in response to environmental signals and regulate polygene quantitative traits. So, TFs would be used to regulate maize PUE. So far, two TFs related to Pi starvation were studied most detailed, which are AtPHRl in Arabidopsis and OsPHR2in rice. They belong to the MYB-CC TF family and are evolutionary homology. Both AtPHRl and OsPHR2were verified to be important regulators for phosphate starvation response and play important roles in the regulation of plant phosphorus accumulation. However, little is known about the functions of PHR-like TFs in maize.
In this study, a homolog gene sequence of AtPHRl was isolated from an inbred line of maize with high PUE. The charicaterization and functions the gene were studied. The results are summarized as follows:
1. Thirty-six maize materials were treated by low phosphorus stress. The absolute values and relative values of dry matter amount and phosphorus accumulate amount were used to be an evaluating indicator to screen high PUE genotypes, which are assisted with analysis of physiological-biochemical charicteristic. Two high PUE genotypes478and TX113were obtained.
2. According to the sequence alignment analysis, a homolog gene sequence of AtPHR1was isolated from an inbred line478of maize with high PUE. Bioinformatics analysis showed that the sequence encodes a449-amino acid protein. Multiple sequence alignment results showed that the predicted amino acids sequence shared43%,64%, and45%identity with the amino acid sequences of AtPHR1, OsPHRl and OsPHR2, respectively. The results also indicated that the four amino acid sequences were highly conserved in both MYB DNA-binding domain (BD) and CC domain. So, the sequence designated as ZmPHRl. Phylogenetic analysis showed that ZmPHRl was clustered in the same group as AtPHRl, OsPHR1and OsPHR2. ZmPHRl was sub-grouped with OsPHR1and OsPHR2, which was consistent with their close evolutionary relationship. ZmPHR1was located on long arm of maize chromosome1. The gene structure of ZmPHR1had a similar splicing pattern as AtPHRl, OsPHRl with seven exons and six introns.
3. The construct with ZmPHR1::GFP fusion protein was introduced into onion epidermal cells by particle bombardment. The fluorescence signal was located predominately in the nuclei. The results demonstrated that ZmPHR1is a nuclear protein and is a MYB-CC TF.ZmPHR1showed a tissue specificity expression pattern, with the highest expression in leaves, the higher expression in roots and the weakest expression in stems. Further qPCR analysis showed that, ZmPHRl expression of different tissues was in a dynamic change along with the prolonging of phosphate starvation treatment time, and exist a positive and negative feedback to phosphate starvation.
4. A binary vector containing the ZmPHRl::GFP fusion protein was constructed and introduced into Arabidopsis by floral dip method. The transgenic Arabidopsis lines with overexpressing ZmPHR1were obtained and verified by Hygromycin resistant screening, DNA-PCR, RT-PCR and GFP.
5. The phenotype of ZmPHRl overexpression lines and wild type (WT) Arabidopsis analysis showed that overexpression of ZmPHRl could significantly improve the growth of transgenic plants. The dry matter yield and Pi content of ZmPHR1-overexpressing lines were much higher than those of WT, especially under LP conditions.
6. The expression patterns of Pi-starvation-inducible genes were examined in ZmPHR1-overexpressing Arabidopsis plants. Under LP conditions, compared to wild-type plants, expression of AtIPS1, AtPS2, AtPS3and AtRNSl in shoots and roots was up-regulated in ZmPHR1overexpression lines, and Pht1;2、Phtl;8and Phtl;9were also up-regulated in roots. It is inferred that the upregulation of these Pi-starvation-inducible genes could increase the plant biomass and phosphorus content.
引文
[1]印莉萍,黄勤妮,吴平.植物营养分子生物学及信号转导.科学出版社(北京)第二版,2006,58-59.
[2]Vance CP, Uhde-Stone C, Allan DL. Phosphorus acquisition and use:critical adaptations by plants for securing a nonrenewable resource. New Phytol,2003,157, 423-447.
[3]徐中平.低磷胁迫对玉米磷吸收、转运及IAA和CTK水平与分布的影响.博士学位论文,2008.
[4]Mimura T. Regulation of phosphate transport and homeostasis in plant cells. Inter Rev Cytol,1999,191,149-200.
[5]Mengel K. Dynamics and availability of major nutrients in soils. Adv Soil Sci,1985, 2,96-100.
[6]Bieleski RL. Phosphate pools, phosphate transport, and phosphate availability. Annu Rev Plant Physiol,1973,24,225-252.
[7]Schachtman DP, Reid RJ, Ayling S M. Phosphorus uptake by plants, from soil to cell. Plant Physiol,1998,116; 447-453.
[8]Raghothama KG, Karthikeyan AS. Phosphate acquisition. Plant Soil,2005,274, 37-49.
[9]鲁如坤.中国土壤营养元素概况(氮、磷、钾).土壤学报,1989,26(3),280-286.
[10]Cordell D, Drangert JO, White S. The story of phosphorus, global food security and food for thought. Glob Environ Change.2009,19,292-305.
[11]Steen I. Phosphorus availability in the 21st century, management of a non-renewable resource. Phosphorus Potassium,1998,217,25-31.
[12]李生秀.植物营养与肥料学科的现状与展望.植物营养与肥料学报,1999,5(3),193-205
[13]王庆仁,李继云,李振声.植物高效利用土壤难溶态磷的研究动态及展望.植物营养与肥料学报,1 998,4(2),107-116.
[14]Holford ICR. Soil phosphorus, its measurement, and its uptake by plants. Aust. J. Soil Res,1997,35,227-239.
[15]Kirkby EA, Johnston AE. Soil and fertilizer phosphorus in relation to crop nutrition. In P J White, JP Hammond, eds, Ecophysiol. of Plant-Phosphorus Inter, Springer Science & Business Media, New York,2008,177-223.
[16]White PJ, Hammond JP. Phosphorus nutrition of terrestrial plant. In PJ White, JP Hammond, eds, Ecophysiol. of Plant-Phosphorus Inter. Springer Science & Business Media, New York,2008,51-81.
[17]Epstein E. Better crops for food, Ciba Foundation Symposium,1983,97(2),61-68.
[18]Marschner H, Romheld V, Horst WJ, et al. Root induced changes in the rhizosphere, importance for mineral nutrition of plants. Zeitschrift fur Pflanzenernachrhung und Bodenkunde,1986,149,441-456.
[19]Gilroy S, Jones DL. Through form to function, root hair development and nutrient uptake. Trends in Plant Sci,2000,5,56-60.
[20]Duff SMG, Sarath G, Plaxton WC. The role of acid phosphatases in plant phosphorus metabolism. Physiol Plant,1994,90,791-800.
[21]Marschner H, Dell B. Nutrient uptake in mycorrhizal symbiosis. Plant Soil,1994, 159,89-102.
[22]Plaxton WC, Carswell MC. Metabolic aspects of the phosphate starvation response in plants. In Plant Responses to Environmental Stresses, From Phytohormones to Genome Reorganization, ed. Lerner, H. R.1999,349-372.
[23]Raghothama KG. Phosphate acquisition. Annu Rev Plant Physiol Plant Mol Biol, 1999,50,665-693.
[24]Lynch JP, Ho MD. Rhizoeconomics, carbon costs of phosphorus acquisition. Plant Soil,2005,269,45-56.
[25]Uhde-Stone C, Gilbert G, Johnson JMF, et al. Acclimation of white lupin to phosphorus deficiency involves enhanced expression of genes related to organic acid metabolism. Plant Soil,2003,248,99-116.
[26]Uhde-Stone C, Zinn KE, Ramirez-Yanez M, et al. Nylon filter arrays reveal differential gene expression in proteoid roots of white lupin in response to P deficiency. Plant Physiol,2003,131(3),1064-1079.
[27]Rychter AM, Mikulska M. The relationship between phosphate status and cyanide-resistant respiration in bean roots. Physiol Plant,1990,79,663-667.
[28]Rychter AM, Chauveau M, Bomsel JL, et al. The effect of phosphate deficiency on mitochondrial activity and adenylate levels in bean roots. Physiol Plant,2006,84, 80-86.
[29]Theodorou ME, Plaxton WC. Metabolic adaptations of plant respiration to nutritional phosphate deprivation. Plant Physiol,1993,101,339-344.
[30]Williamson LC, Ribrioux SPCP, Fitter AH, et al. Phosphate availability regulates root system architecture in Arabidopsis. Plant Physiol,2001,126,875-882.
[31]Lopez-Bucio J, Cruz-Ramirez A, Herrera-Estrella L. The role of nutrient availability in regulating root architecture. Curr Opin Plant Biol,2003,6,280-287.
[32]Lambers HY, Shane MW, Cramer MD, et al. Root structure and functioning for efficient acquisition of phosphorus, matching morphological and physiological traits. Ann Bot,2006,98,693-713.
[33]Bates TR, Lynch JP. Stimulation of root hair elongation in Arabidopsis thaliana by low phosphorus availability. Plant Cell Environ,1996,19,529-538.
[34]Watts M, Evans JR. Proteoid roots Physiology and development. Plant Physiol, 1999,121,317-323.
[35]Lynch JP, Brown KM. Topsoil foraging-an architectural adaptation of plants to low phosphorus. Plant Soil,2001,237,225-237.
[36]Lynch PJ. Root architecture and plant productivity. Plant Physiol,1995,109, 7-13.
[37]Burleigh SH, Harrison MJ. A novel gene whose expression in Medicago truncatula roots is suppressed in response to colonization by vesicular-arbuscular mycorrhizal (VAM) fungi and to phosphate nutrition. Plant Mol Biol,1997,34,199-208.
[38]Harrison MJ. Molecular and cellular aspects of the arbuscular mycorrhizal symbiosis. Plant Mol Biol,1999,50,219-243.
[39]Lopez-Bucio J, de la Vega OM, Guevara-Garcia A, et al. Enhanced phosphorus uptake in transgenic tobacco plants that overproduce citrate. Nat Biotechnol,2000, 18,450-453.
[40]Jones DL. Organic acids in the rhizosphere, a critical review. Plant Soil,1998,205, 25-44.
[41]李继云,等.有效利用土壤营养元素的作物育种新技术研究中国科学B辑.1995,25(1),41-48.
[42]Dinkelaker B, Romheld V, Marschner H. Citric acid excretion and precipitation of calcium citrate in the rhizosphere of white lupin (Lupinus albus L.). Plant, Cell Environ,1989,12,285-292.
[43]李德华,向春雷,姜益泉,等.低磷胁迫下水稻不同品种根系有机酸分泌的差异.中国农学通报,2005,21(11),186-188.
[44]王树起,韩晓增,乔云发,等.缺磷胁迫条件下大豆根系有机酸的分泌特性.大豆科学,2009,28(3),409-414.
[45]Reid MS, Bieleski RL. Changes in phosphatase activity in phosphorus-deficient spirodda. Planta,1970,94,273-281.
[46]Goldstein AH, Baertlein DA, McDaniel RG. Phosphate starvation inducible metabolism in Lycopersicon esculentum Ⅰ. Excretion of acid phosphatase by tomato plants and suspension-cultured cells. Plant Physiol,1988,87,711-715.
[47]Goldstein AH, Danon A, Baertlein DA, et al. Phosphate starvation inducible metabolism in Lycopersicon esculentum Ⅱ Characterization of the psi excreted acid phosphatase. Plant Physiol,1988,87,716-720.
[48]Helal HM. Varietal differences in root phosphatase activity as related to the utilization of organic phosphates. Plant and Soil,1990,123,161-163.
[49]Bozzo GG, Raghothama KG. Plaxton WC. Structural and kinetic properties of a novel purple acid phosphatase from phosphate-starved tomato (Lycopersicon esculentum) cell cultures. Biochem J,2004,377,419-428.
[50]McLachlan K, Elliott D, DeMarco D, et al. Leaf acid phosphatase isozymes in the diagnosis of phosphorus status in field-grown wheat. Aust J Agric Res,1987,38, 1-13.
[51]张海伟,黄宇,叶祥盛,等.低磷胁迫下甘蓝型油菜酸性磷酸酶对磷效率的贡献分析.中国科学(生命科学),2010,40(5),418-427.
[52]杜育梅,白昌军,田 江,等.柱花草适应酸性缺磷土壤的基因型差异及可能的生理机制.华南农业大学学报,2008,29(4),6-11.
[53]Xu QZ. Acid phosphatase act ivity in wheat leaves and effects of phosphorus def iciency. Plant Physiol Comm,1989,6,20-23.
[54]Yan X, Liao H, Trull MC, et al. Induction of a major leaf acid phosphatase does not confer adaptation to low phosphorus availability in common bean. Plant Physiol, 2001,125,1901-1911.
[55]张斌,秦岭.植物对低磷胁迫的适应及其分子基础.分子植物育种,2010,8(4),776-783.
[56]Nurnberger T, Abel S, Jost W, et al. Induction of an extracellular ribonuclease in cultured tomato cells upon phosphate starvation. Plant Physiol,1990,92,970-976.
[57]Loffler A, Abel S, Jost W, et al. Phosphate-regulated induction of intracellular ribonucleases in cultured tomato (Lycopersicon esculentum) cells. Plant Physiol, 1992,98,1472-1478.
[58]Goldstein AH, Baertlein DA, Danon A. Phosphate starvation stress as an experimental system for molecular analysis. Plant Mol Biol Rep,1989,7(1),7-16.
[59]Bariola PA, Howard CJ, Taylort CB, et al. The Arabidopsis ribonuclease gene RNS1 is tightly controlled in response to phosphate limitation. Plant J,1994,6(5), 673-685.
[60]Morcuende R, Bari R, Gibon Y, et al. Genome-wide reprogramming of metabolism and regulatory networks of Arabidopsis in response to phosphorus. Plant Cell Environ,2007,30,85-112.
[61]Dodds PN, Clarke AE, Newbigin E. Molecular characterization of an S-like RNase of Nicotiana alata that is induced by phosphate starvation. Plant Mol. Biol,1996, 31,227-238.
[62]Kock M, Theierl K, Stenzel I, et al. Extracellular administration of phosphate-sequestering metabolites induces ribonucleases in cultured tomato cells. Planta,1998,204,404-407.
[63]Poirier Y, Bucher M. Phosphate transport and homeostasis in Arabidopsis, In, Chris S, Elliot M, Jeff D, et al. The Arabidopsis Book, Am Soc Plant Bio,2002, 1-35.
[64]Muchhal US, Pardot JM, Raghothama KG. Phosphate transporters from the higher plant Arabidopsis thaliana. Proc. Natl. Acad. Sci. USA,1996,9(93),10519-10523.
[65]Daram P, Brunner S, Rausch C, et al. Pht2;l encodes a low-affinity phosphate transporter from Arabidopsis. Plant Cell,1999,11,2153-2166.
[66]Versaw WK, Harrison MJ. A Chloroplast phosphate transporter, PHT2;1, influences allocation of phosphate within the plant and phosphate-starvation responses. Plant Cell,2002,14,1751-1766.
[67]Rausch C, Zimmermann P, Amrhein N, et al. Expression analysis suggests novel roles for the plastidic phosphate transporter Pht2;l in auto-and heterotrophic tissues in potato and Arabidopsis. Plant J,2004,39(1),13-28.
[68]Ticconi CA, Abel S. Short on phosphate, plant surveillance and countermeasures, Trends Plant Sci,2004,9(11),548-555.
[69]Nandi SK, Pant RC, Nissen P. Multiphasic uptake of phosphate by corn roots. Plant, Cell Environ,1987,10,463-474.
[70]Furihata T, Suzuki M, Sakurai H. Kinetic characterization of two phosphate uptake systems with different affinities in suspension-cultured Catharanthus roseus protoplasts. Plant, Cell Physiol,1992,33,1151-1157.
[71]Daram P, Brunner S, Persson BL, et al. Functional analysis and cell-specific expression of a phosphate transporter from tomato. Planta,1998,206,225-233.
[72]Liu H, Trieu AT, Blaylock LA. Cloning and characterization of two phosphate transporters from Medicago truncatula roots, regulation in response to phosphate and to colonization by arbuscular mycorrhizal fungi. Mol Plant Micro Interact, 1998,11,14-22.
[73]Nussaume L, Kanno S, Javot H, et al. Phosphate import in plants, focus on the PHT1 transporters. Front Plant Sci,2011,2,1-12.
[74]Paszkowski U, Kroken S, Roux C, et al. Rice phosphate transporters include an evolutionarily divergent gene specifically activated in arbuscular mycorrhizal symbiosis. Proc Natl Acad Sci,2002,99,13324-13329.
[75]Ai PH, Sun SB, Zhao JN, et al. Two rice phosphate transporters, OsPht1;2 and OsPht1;6 have different functions and kinetic properties in uptake and translocation. Plant J,2009,57,798-809.
[76]Jia HF, Ren HY, Gu M, et al. The phosphate transporter gene OsPhtl;8 is involved in phosphate homeostasis in rice. Plant Physiol,2011,156,1164-1175.
[77]Smith FW, Cybinski DH, Rae AL. Regulations of expression of genes encoding phosphate transporters in barley roots. Dordrecht, Kluwer Academic Publishers, 1999.
[78]Rae AL, Cybinski DH, Jarmey JM, et al. Characterization of two phosphate transporters from barley, evidence for diverse function and kinetic properties among members of the Phtl family. Plant Mol Biol,2003,53,27-36.
[79]Schunmann PHD, Richardson AE, Smith FE, et al. Characterization of promoter expression patterns derived from the Phtl phosphate transporter genes of barley (Hordeum vulgare L.). JExp Bot,2004,55(399),855-865.
[80]Nagy R, Vasconcelos MJ, Zhao S, et al. Differential regulation of five Phtl phosphate transporters from maize (Zea mays L.). Plant Biol,2006,8,186-197.
[81]Leggewie G, Willmitzer L, Riesmeier J W. Two cDNAs from potato are able to complement a phosphate uptake-deficient yeast mutant, identification of phosphate transporters from higher plants. Plant Cell,1997,9,381-392.
[82]Liu C, Muchhal US, Uthappa M, et al. Tomato phosphate transporter genes are differentially regulated in plant tissues by phosphorus. Plant Physiol,1998 a,116, 91-99.
[83]Nagy R, Karandashov V, Chague V, et al. The characterization of novel mycorrhiza-specific phosphate transporters from Lycopersicon esculentum and Solanum tuberosum uncovers functional redundancy in symbiotic phosphate transport in Solanaceous species. Plant J,2005,42,236-250.
[84]Xu GH, Chague V, Melamed-Bessud C, et al. Isolation and functional characterization of LePT4, a mycorrhizal enhanced phosphate transporter in tomato, In, Li C J, Zhang F S, Doberman A. (eds.). Plant Nut Food Sec, Human Health Enviro Prot, Proc of 15th Inter Plant Nut Col, Tsinghua University Press, Beijing, China,2005,90-91.
[85]Baek SH, Chung IM, Yun SJ. Molecular cloning and characterization of a tobacco leaf cDNA encoding a phosphate transporter. Mol Cell,2001,11,1-6.
[86]Harrison MJ, Dewbre GR, Liu J. A phosphate transporter from Medicago truncatula involved in the acquisition of phosphate released by arbuscular mycorrhizal fungi. Plant Cell,2002,14,2413-2429.
[87]Javot H, Penmetsa RV, Terzaghi N, et al. A Medicago truncatula phosphate transporter indispensable for the arbuscular mycorrhizal symbiosis. Proc Natl Acad Sci,2007,104(5),1720-1725.
[88]常胜合,舒海燕,童一平,等.两个小麦磷转运蛋白基因的分离、功能鉴定和表 达研究.西北植物学报,2004,24(10),1779-1785.
[89]Davies TGE, Ying J, Xu Q, et al. Expression analysis of putative high-affinity phosphate transporters in Chinese winter wheats. Plant Cell Environ,2002,5, 1325-1339.
[90]Liu JQ, Uhde-Stone C, Li AG, et al. A phosphate transporter with enhanced expression in proteoid roots of white lupin (Lupinus albus L.). Plant Soil,2001, 237,257-266.
[91]Yang CY, Wang YX, Shen H, et al. Cloning and characterizaton of a novel root-specific putative phosphate transporter in soybean (Glycine max L.), In, Li C J, Zhang FS, Doberman A. (eds.). Plant Nut Food Sec, Human Health Enviro Prot, Proc 15th Inter Plant Nut Col, Tsinghua University Press, Beijing, China,2005, 172-173.
[92]Kai M, Masuda Y, Kikuchi Y, et al. Isolation and characterization of a cDNA from Catharanthusroseus which is highly homologous with phosphate transporter. Soil Sci Plant Nutr,1997,43,227-235.
[93]Loth-Pereda V, Orsini E, Courty PE, et al. Structure and expression profile of the phosphate Phtl transporter gene family in mycorrhizal Populustrichocarpa. Plant Physiol,2011,156,2141-2154.
[94]Mudge SR, Rae AL, Diatloff E, et al. Expression analysis suggests novel roles for members of the Phtl family of phosphate transporters in Arabidopsis. Plant J, 2002,31(3),341-353.
[95]Rausch C, Daram P, Brunner S, et al. A phosphate transporter expressed in arbuscular-containing cells in potato. Nature,2001,414,462-466.
[96]李锋,李木英,潘晓华等.不同水稻品种幼苗适应低磷胁迫的根系生理生化特性.中国水稻科学,2004,18(1),48-52.
[97]Hammond JP, Bennett MJ, Bowen HC, et al. Changes in gene expression in Arabidopsis shoots during phosphate starvation and the potential for developing smart plants. Plant Physiol,2003,132,578-596.
[98]Misson J, Raghothama KG, Jain A, et al. A genome-wide transcriptional analysis using Arabidopsis thaliana Affymetrix gene chips determined plant responses to phosphate deprivation. Proc Natl Acad Sci,2005,102,11934-11939.
[99]Wasaki J, Yonetani R, Kuroda S, et al. Transcriptomic analysis of metabolic changes by phosphorus stress in rice plant roots. Plant, Cell Environ,2003,26, 1515-1523.
[100]Wasaki J, Yonetani R, Shinano T, et al. Expression of the OsPIl gene, cloned from rice roots using cDNA microarray, rapidly responds to phosphorus status. New Phytol,2003,158,239-248.
[101]Tian J, Venkatachalam P, Liao H, et al. Molecular cloning and characterization of phosphorus starvation responsive genes in common bean (Phaseolus vulgaris L.). Planta,2007,227,151-165.
[102]Calderon-Vazquez C, Ibarra-Laclette E, Caballero-Perez J, et al. Transcript profiling of Zea mays roots reveals gene responses to phosphate deficiency at the plant-and species-specific levels. J Exp Botany,2008,59(9),2479-2497.
[103]Calderon-Vazquez C, Sawers RJH, Herrera-Estrella L. Phosphate deprivation in maize, genetics and genomics. Plant Physiol,2011,156,1067-1077.
[104]Wu P, Ma L, Hou XL, et al. Phosphate starvation triggers distinct alterations of genome expression in srabidopsis roots and leaves. Plant Physiol,2003,132, 1260-1271.
[105]Li LH, Liu C, Lian XM. Gene expression profiles in rice roots under low phosphorus stress. Plant Mol Biol,2010,72,423-432.
[106]Cai HM, Xie WB, Zhu T, et al. Transcriptome response to phosphorus starvation in rice. Acta Physiol Plant,2012,34,327-341.
[107]Gallou A, Declerck S, Cranenbrouck S. Transcriptional regulation of defence genes and involvement of the WRKY transcription factor in arbuscular mycorrhizal potato root colonization. Funct Integr Genomics,2012,12,183-198.
[108]Ren F, Chen L, Guo QQ, et al. Identification and expression analysis of genes induced by phosphate starvation in leaves and roots of Brassica napus. Plant Growth Regul,2011,65,65-81.
[109]Haran S, Logendra S, Seskar M, et al. Characterization of Arabidopsis acid phosphatase promoter and regulation of acid phosphatase expression. Plant Physiol,2000,124,615-626.
[110]Burleigh SH, Harrison MJ. The down-regulation of Mt4-like genes by phosphate fertilization occurs systemically and involves phosphate translocation to the shoots. Plant Physiol,1999,119,241-248.
[111]Liu C, Muchhal US, Raghothama KG. Differential expression of TPS11, a phosphate starvation-induced gene in tomato. Plant Mol Biol,1997,33,867-874.
[112]Martin AC, Del Pozo JC, Iglesias J, et al. Influence of cytokinins on the expression of phosphate starvation responsive genes in Arabidopsis. Plant J,2000, 24,559-567.
[113]Muchhal US, Raghotama KG. Transcriptional regulation of plant phosphate transporters. Proc Natl Acad Sci,1999,96,5868-5872.
[114]Mitsukawa N, Okumura S, Shirano Y, et al. Overexpression of an Arabidopsis thaliana high-affinity phosphate transporter gene in tobacco cultured cells enhances cell growth under phosphate-limited conditions. Proc Natl Acad Sci, 1997,94,7098-7102.
[115]Park MR, Baek SH, De los Reyes BG, et al. Overexpression of a high-affinity phosphate transporter gene from tobacco (NtPTI) enhances phosphate uptake and accumulation in transgenic rice plants. Plant Soil,2007,292,259-269.
[116]Coelho GTCP, Carneiro NP, Karthikeyan AS, et al. A phosphate transporter promoter from Arabidopsis thaliana AtPHT1;4 gene drives preferential gene expression in transgenic maize roots under phosphorus starvation. Plant Mol Biol Rep,2010,28,717-723.
[117]Shin H, Shin HS, Dewbre GR, et al. Phosphate transport in Arabidopsis, Phtl;1 and Phtl;4 play a major role in phosphate acquisition from both low-and high-phosphate environments. Plant J,2004,39,629-642.
[118]Nagarajan VK, Jain A, Poling MD, et al. Arabidopsis Phtl;5 mobilizes phosphate between source and sink organs and influences the interaction between phosphate homeostasis and ethylene signaling. Plant Physiol,2011,156,1149-1163.
[119]Liu F, Wang ZY, Ren HY, et al. OsSPX1 suppresses the function of OsPHR2 in the regulation of expression of OsPT2 and phosphate homeostasis in shoots of rice. Plant J,2010,62,508-517.
[120]Sun SB, Gu M, Cao Y, et al. A constitutive expressed phosphate transporter, OsPhtl;1, modulates phosphate uptake and translocation in phosphate-replete rice. Plant Physiol,2012,8(159),1571-1581.
[121]Baldwin JC, Karthikeyan AS, Cao AQ, et al. Biochemical and molecular analysis of LePS2;l, a phosphate starvation induced protein phosphatase gene from tomato. Planta,2008,228,273-280.
[122]Baldwin JC, Karthikeyan AS, Raghothama KG. LEPS2, a phosphorus starvation-induced novel acid phosphatase from tomato. Plant Physiol,2001,125, 728-737.
[123]Martin T, Hellmann H, Schmidt R, et al. Identification of mutants in metabolically regulated gene expression. Plant J.1997,11,53 62.
[124]Hou X L, Wu P, Jiao FC, et al. Regulation of the expression of OsIPSl and OsIPS2 in rice via systemic and local Pi signalling and hormones. Plant, Cell Environ. 2005,28,353 364.
[125]Muller R, Morant M, Jarmer H, et al. Genome-wide analysis of the Arabidopsis leaf transcriptome reveals interaction of phosphate and sugar metabolism. Plant Physiol,2001,143,156-171.
[126]Rubio V, Linhares F, Solano R, et al. A conserved MYB transcription factor involved in phosphate starvation signaling both in vascular plants and in unicellular algae. Genes Dev,2001,15,2122-2133.
[127]Nilsson L, Muller R, Nielsen TH. Increased expression of the MYB-related transcription factor, PHR1, leads to enhanced phosphate uptake in Arabidopsis thaliana. Plant, Cell Environ,2007,30,1499-1512.
[128]Devaiah BN, Karthikeyan AS, Raghothama KG. WRKY75 transcription factor is a modulator of phosphate acquisition and root development in Arabidopsis. Plant Physiol,2001,143,1789-1801.
[129]Devaiah BN, Nagarajan VK, Raghothama KG. Phosphate homeostasis and root development in Arabidopsis is synchronized by the zinc finger transcription factor ZAT6. Plant Physiol,2007,145,147-159.
[130]Devaiah BN, Madhuvanthi R, Karthikeyan AS, et al. Phosphate starvation responses and gibberellic acid biosynthesis are regulated by the MYB62 transcription factor in Arabidopsis. Mol Plant,2009,2,43-58.
[131]Chen YF, Li LQ, Xu Q, et al. The WRKY6 transcription factor modulates PHOSPHATE1 expression in response to low Pi stress in Arabidopsis. Plant Cell, 2009,21,3554-3566.
[132]Chen ZH, Nimmo GA, Jenkins GI, et al. BHLH32 modulates several biochemical and morphological processes that respond to Pi starvation in Arabidopsis. Biochem J,2007,405,191-198.
[133]Yi KK, Wu Z, Zhou J, et al. OsPTFl, a novel transcription factor involved in tolerance to phosphate starvation in rice. Plant Physiol,2005,138,2087-2096.
[134]Tang Z, Sadka A, Morishige D T, et al. Homeodomain leucine zipper proteins bind to the phosphate response domain of the soybean VspB tripartite promoter. Plant Physiol,2001,125,797-809.
[135]Wykoff DD, Grossman AR, Weeks DP, et al. Psrl, a nuclear localized protein that regulates phosphorus metabolism in Chlamydomonas. Proc Natl Acad Sci,1999, 96,15336-15341.
[136]Franco-Zorrilla JM, Gonzalez E, Bustos R, et al. The transcriptional control of plant responses to phosphate limitation. JExp Botany,2004,55(396),285-293,
[137]Schunmann PHD, Richardson AE, Vickers C E, et al. Promoter analysis of the barley Phtl;l phosphate transporter gene identifies regions controlling root expression and responsiveness to phosphate deprivation. Plant Physiol,2004,136, 4205-4214.
[138]Del Pozo JC, Allona I, Rubio V, et al. A type 5 acid phosphatase gene from Arabidopsis thaliana is induced by phosphate starvation and by some other types of phosphate mobilizing/oxidative stress conditions. Plant J,1999,19,579-589.
[139]Li D, Zhu H, Liu K, et al. Purple acid phosphatases of Arabidopsis thaliana, comparative analysis and differential regulation by phosphate deprivation. J Biol Chem,2002,277,27772-27781.
[140]Okumura S, Mitsukawa N, Shirano Y, et al. Phosphate transporter gene family of Arabidopsis thaliana. DNA Res,1998,5,261-269.
[141]Mudge SR, Rae AL, Diatloff E, et al. Expression analysis suggests novel roles for members of the Phtl family of phosphate transporters in Arabidopsis. Plant J, 2002,31,341-353.
[142]Taylor CB, Bariola PA, del Cardayre SB, et al. RNS2, a senescence-associated RNase of Arabidopsis that diverged from the S-RNases before speciation. Proc Natl Acad Sci,1993,90,5118-5122.
[143]Schachtman DP, Shin R. Nutrient sensing and signaling. Annu Rev Plant Biol,2007, 58,47-69.
[144]Miura K, Rus A, Sharkhuu A, et al. The Arabidopsis SUMO E3 ligase SIZ1 controls phosphate deficiency responses. Proc Natl Acad Sci,2005,102, 7760-7765.
[145]Bari R, Pant BD, Stitt M, et al. PHO2, microRNA399, and PHR1 define a phosphate-signaling pathway in plants. Plant Physiol,2006,141,988-999.
[146]Pant BD, Buhtz A, Kehr J, et al. MicroRNA399 is a long-distance signal for the regulation of plant phosphate homeostasis. Plant J,2008,53,731-773.
[147]Valli A, Todesco M, Mateos I, et al. Target mimicry provides a new mechanism for regulation of microRNA activity. Nat Genet,2007,39,1033-1037.
[148]Chiou TJ, Aung K, Lin SI, et al. Regulation of phosphate homeostasis by microRNA in Arabidopsis. Plant Cell,2006,18,412-421.
[149]Aung K, Lin SI, Wu CC, et al. Pho2, a phosphate overaccumulator, is caused by a nonsense mutation in a microRNA399 target gene. Plant Physiol,2006,141, 1000-1011.
[150]Fujii H, Chiou TJ, Lin SI, et al. A miRNA involved in phosphate-starvation response in Arabidopsis. Current Biol,2005,15,2038-2043.
[151]Lin SI, Chiang SF, Lin WY, et al. Regulatory network of microRNA399 and PHO2 by systemic signaling. Plant Physiol,2008,147,732-746.
[152]Franco-Zorrilla JM, Valli A, Todesco M, et al. Target mimicry provides a new mechanism for regulation of microRNA activity. Nat Genet,2007,39,1033-1037.
[153]Gonzalez E, Solano R, Rubio V, et al. Phosphate Transporter Traffic Facilitator 1 is a plant-specific SEC12-related protein that enables the endoplasmic reticulum exit of a high affinity phosphate transporter in Arabidopsis. Plant Cell.2005,17, 3500-3512.
[154]Zhou J, Jiao FC, Wu ZC, et al. OsPHR2 is involved in phosphate-starvation signaling and excessive phosphate accumulation in shoots of plants. Plant Physiol, 2008,146,1673-1686.
[155]Wu P, Wang XM. Role of OsPHR2 on phosphorus homeostasis and root hairs development in rice (Oryza sativa L.). Plant Signal Beh,2008,3,674-675.
[156]Kaeppler SM, Parke JL, Mueller SM, et al. Variation among maize inbred lines and detection of quantitative trait loci for growth at low phosphorus and responsiveness to Arbuscular mycorrhizal fungi. Crop Sci,2000,40,358-363.
[157]Da Silva AE, Gabelman W. Screening maize inbred lines for tolerance to low-P stress condition. Plant Soil,1992,146,181-187.
[158]张吉海,高世斌,杨克诚,等.玉米耐低磷种质资源的筛选与鉴定.植物遗传资源学报,2008,9(3),335-339.
[159]张丽梅,贺立源,李建生,等.玉米自交系耐低磷材料苗期筛选研究.中-国农业科学,2004,37(12),1955-1959.
[160]Vance CP. Quantitative trait loci, epigenetics, sugars, and microRNAs: Quaternaries in phosphate acquisition and use. Plant Physiol,2010,154,582 588.
[161]Century K, Lynne RT, Ratcliffe O J. Regulating the regulators, the future prospects for transcription-factor-based agricultural biotechnology products. Plant physiol, 2008,147(1),20-29.
[162]明凤,米国华,张福锁,等.水稻对低磷反应的基因型差异及其生理适应机制的初步研究.应用与环境生物学报,2000,6(2),1 38-141.
[163]华瑞,沈玉芳,李世清,等.小麦/玉米苗期磷累积量对介质供磷水平反应的差异.中国生态农业学报,2009,1 7(3),429-435.
[164]孔忠新,杨丽丽,张政值,等.小麦耐低磷基因型的筛选.麦类作物学报,201 0,30(4),591-595.
[165]丁洪,李生秀.大豆品种耐低磷和对磷肥效应的遗传差异.植物营养与肥料学报,1998,4(3),257-263.
[166]Lynch J. The role of nutrient-efficient crops in modern agriculture. J Crop Prod, 1998,1(2),241-264.
[167]Barry DAJ, Miller MH. Phosphorus nutritional requirement of maize seedlings for maximum yield. Agron J,1989,81,95-99.
[168]李绍长,龚江,王军.玉米自交系苗期耐低磷基因型的筛选.玉米科学,2003,11(3),85-89.
[169]吴平,倪俊健.应用AFLP与RFLP标记研究水稻磷吸收与利用率的数量性 状位点.植物学报,2000,42(3),229-233.
[170]龚江,李绍长,夏春兰,等.低磷胁迫下玉米自交系磷高效基因型筛选.新疆农业科学,2002,39(2),77-81.
[171]杜育梅,白昌军,田江,等.柱花草适应酸性缺磷土壤的基因型差异及可能的生理机制.华南农业大学学报,2008,29(4),6-11.
[172]章爱群,贺立源,门玉英,等.磷水平对不同耐低磷玉米基因型幼苗生长和养分吸收的影响.应用与环境生物学报,2008,14(3),347-350.
[173]Martin C, Paz-Ares J. MYB transcription factors in plants. Trends Genet,1997,13, 67-73.
[174]刘蕾,杜海,唐晓凤,等.MYB转录因子在植物抗逆胁迫中的作用及其分子机理,遗传,2008,30(10),1265-1271.
[175]Livak KJ, Schmittgen TD. Analysis of relative gene expression data using real-time quantitative PCR and the 2(-Delta Delta C(T)). Methods,2001,25,402-408.
[176]Clough SJ, Bent AF. Floral dip, a simplified method for Agrobacterium-mediated transformation of Arabidopsis thaliana. Plant J,1998,16,735 743.
[177]黄国存,朱生伟,董越梅,等.绿色荧光蛋白及其在植物研究中的应用.植物学通报,1998,15(5),24-30.
[178]Alexander M, Markus I, Dieter H. Stable transformation of sunflower(Helianthus annuus L.) using a non-meristematic regeneration protocol and green fluorescent protein as a vital marker. Trans Res,2001,10,435 444.
[179]Inger BH, Henrik BP, Mette, et al. Transformation of barley (Hordeum vulgare L.) by Agrobacterium tumefaciens infection of in vitro cultured ovules. Plant Cell Rep, 2006,25,1325 1335.
[180]Abel S, Ticconi CA, Delatorre, CA. Phosphate sensing in higher plants. Physiol. Plant,2002,115,1-8.
[181]May A, Berger S, Hertel T, et al. The Arabidopsis thaliana phosphate starvation responsive gene AtPPsPasel encodes a novel type of inorganic pyrophosphatase. Biochim Biophys Acta,2011,1810,178 185.
[182]Raghothama KG. Phosphate transport and signaling. Curr Opin Plant Biol,2000,3, 182 187.
[2]Vance CP, Uhde-Stone C, Allan DL. Phosphorus acquisition and use:critical adaptations by plants for securing a nonrenewable resource. New Phytol,2003,157, 423-447.
[3]徐中平.低磷胁迫对玉米磷吸收、转运及IAA和CTK水平与分布的影响.博士学位论文,2008.
[4]Mimura T. Regulation of phosphate transport and homeostasis in plant cells. Inter Rev Cytol,1999,191,149-200.
[5]Mengel K. Dynamics and availability of major nutrients in soils. Adv Soil Sci,1985, 2,96-100.
[6]Bieleski RL. Phosphate pools, phosphate transport, and phosphate availability. Annu Rev Plant Physiol,1973,24,225-252.
[7]Schachtman DP, Reid RJ, Ayling S M. Phosphorus uptake by plants, from soil to cell. Plant Physiol,1998,116; 447-453.
[8]Raghothama KG, Karthikeyan AS. Phosphate acquisition. Plant Soil,2005,274, 37-49.
[9]鲁如坤.中国土壤营养元素概况(氮、磷、钾).土壤学报,1989,26(3),280-286.
[10]Cordell D, Drangert JO, White S. The story of phosphorus, global food security and food for thought. Glob Environ Change.2009,19,292-305.
[11]Steen I. Phosphorus availability in the 21st century, management of a non-renewable resource. Phosphorus Potassium,1998,217,25-31.
[12]李生秀.植物营养与肥料学科的现状与展望.植物营养与肥料学报,1999,5(3),193-205
[13]王庆仁,李继云,李振声.植物高效利用土壤难溶态磷的研究动态及展望.植物营养与肥料学报,1 998,4(2),107-116.
[14]Holford ICR. Soil phosphorus, its measurement, and its uptake by plants. Aust. J. Soil Res,1997,35,227-239.
[15]Kirkby EA, Johnston AE. Soil and fertilizer phosphorus in relation to crop nutrition. In P J White, JP Hammond, eds, Ecophysiol. of Plant-Phosphorus Inter, Springer Science & Business Media, New York,2008,177-223.
[16]White PJ, Hammond JP. Phosphorus nutrition of terrestrial plant. In PJ White, JP Hammond, eds, Ecophysiol. of Plant-Phosphorus Inter. Springer Science & Business Media, New York,2008,51-81.
[17]Epstein E. Better crops for food, Ciba Foundation Symposium,1983,97(2),61-68.
[18]Marschner H, Romheld V, Horst WJ, et al. Root induced changes in the rhizosphere, importance for mineral nutrition of plants. Zeitschrift fur Pflanzenernachrhung und Bodenkunde,1986,149,441-456.
[19]Gilroy S, Jones DL. Through form to function, root hair development and nutrient uptake. Trends in Plant Sci,2000,5,56-60.
[20]Duff SMG, Sarath G, Plaxton WC. The role of acid phosphatases in plant phosphorus metabolism. Physiol Plant,1994,90,791-800.
[21]Marschner H, Dell B. Nutrient uptake in mycorrhizal symbiosis. Plant Soil,1994, 159,89-102.
[22]Plaxton WC, Carswell MC. Metabolic aspects of the phosphate starvation response in plants. In Plant Responses to Environmental Stresses, From Phytohormones to Genome Reorganization, ed. Lerner, H. R.1999,349-372.
[23]Raghothama KG. Phosphate acquisition. Annu Rev Plant Physiol Plant Mol Biol, 1999,50,665-693.
[24]Lynch JP, Ho MD. Rhizoeconomics, carbon costs of phosphorus acquisition. Plant Soil,2005,269,45-56.
[25]Uhde-Stone C, Gilbert G, Johnson JMF, et al. Acclimation of white lupin to phosphorus deficiency involves enhanced expression of genes related to organic acid metabolism. Plant Soil,2003,248,99-116.
[26]Uhde-Stone C, Zinn KE, Ramirez-Yanez M, et al. Nylon filter arrays reveal differential gene expression in proteoid roots of white lupin in response to P deficiency. Plant Physiol,2003,131(3),1064-1079.
[27]Rychter AM, Mikulska M. The relationship between phosphate status and cyanide-resistant respiration in bean roots. Physiol Plant,1990,79,663-667.
[28]Rychter AM, Chauveau M, Bomsel JL, et al. The effect of phosphate deficiency on mitochondrial activity and adenylate levels in bean roots. Physiol Plant,2006,84, 80-86.
[29]Theodorou ME, Plaxton WC. Metabolic adaptations of plant respiration to nutritional phosphate deprivation. Plant Physiol,1993,101,339-344.
[30]Williamson LC, Ribrioux SPCP, Fitter AH, et al. Phosphate availability regulates root system architecture in Arabidopsis. Plant Physiol,2001,126,875-882.
[31]Lopez-Bucio J, Cruz-Ramirez A, Herrera-Estrella L. The role of nutrient availability in regulating root architecture. Curr Opin Plant Biol,2003,6,280-287.
[32]Lambers HY, Shane MW, Cramer MD, et al. Root structure and functioning for efficient acquisition of phosphorus, matching morphological and physiological traits. Ann Bot,2006,98,693-713.
[33]Bates TR, Lynch JP. Stimulation of root hair elongation in Arabidopsis thaliana by low phosphorus availability. Plant Cell Environ,1996,19,529-538.
[34]Watts M, Evans JR. Proteoid roots Physiology and development. Plant Physiol, 1999,121,317-323.
[35]Lynch JP, Brown KM. Topsoil foraging-an architectural adaptation of plants to low phosphorus. Plant Soil,2001,237,225-237.
[36]Lynch PJ. Root architecture and plant productivity. Plant Physiol,1995,109, 7-13.
[37]Burleigh SH, Harrison MJ. A novel gene whose expression in Medicago truncatula roots is suppressed in response to colonization by vesicular-arbuscular mycorrhizal (VAM) fungi and to phosphate nutrition. Plant Mol Biol,1997,34,199-208.
[38]Harrison MJ. Molecular and cellular aspects of the arbuscular mycorrhizal symbiosis. Plant Mol Biol,1999,50,219-243.
[39]Lopez-Bucio J, de la Vega OM, Guevara-Garcia A, et al. Enhanced phosphorus uptake in transgenic tobacco plants that overproduce citrate. Nat Biotechnol,2000, 18,450-453.
[40]Jones DL. Organic acids in the rhizosphere, a critical review. Plant Soil,1998,205, 25-44.
[41]李继云,等.有效利用土壤营养元素的作物育种新技术研究中国科学B辑.1995,25(1),41-48.
[42]Dinkelaker B, Romheld V, Marschner H. Citric acid excretion and precipitation of calcium citrate in the rhizosphere of white lupin (Lupinus albus L.). Plant, Cell Environ,1989,12,285-292.
[43]李德华,向春雷,姜益泉,等.低磷胁迫下水稻不同品种根系有机酸分泌的差异.中国农学通报,2005,21(11),186-188.
[44]王树起,韩晓增,乔云发,等.缺磷胁迫条件下大豆根系有机酸的分泌特性.大豆科学,2009,28(3),409-414.
[45]Reid MS, Bieleski RL. Changes in phosphatase activity in phosphorus-deficient spirodda. Planta,1970,94,273-281.
[46]Goldstein AH, Baertlein DA, McDaniel RG. Phosphate starvation inducible metabolism in Lycopersicon esculentum Ⅰ. Excretion of acid phosphatase by tomato plants and suspension-cultured cells. Plant Physiol,1988,87,711-715.
[47]Goldstein AH, Danon A, Baertlein DA, et al. Phosphate starvation inducible metabolism in Lycopersicon esculentum Ⅱ Characterization of the psi excreted acid phosphatase. Plant Physiol,1988,87,716-720.
[48]Helal HM. Varietal differences in root phosphatase activity as related to the utilization of organic phosphates. Plant and Soil,1990,123,161-163.
[49]Bozzo GG, Raghothama KG. Plaxton WC. Structural and kinetic properties of a novel purple acid phosphatase from phosphate-starved tomato (Lycopersicon esculentum) cell cultures. Biochem J,2004,377,419-428.
[50]McLachlan K, Elliott D, DeMarco D, et al. Leaf acid phosphatase isozymes in the diagnosis of phosphorus status in field-grown wheat. Aust J Agric Res,1987,38, 1-13.
[51]张海伟,黄宇,叶祥盛,等.低磷胁迫下甘蓝型油菜酸性磷酸酶对磷效率的贡献分析.中国科学(生命科学),2010,40(5),418-427.
[52]杜育梅,白昌军,田 江,等.柱花草适应酸性缺磷土壤的基因型差异及可能的生理机制.华南农业大学学报,2008,29(4),6-11.
[53]Xu QZ. Acid phosphatase act ivity in wheat leaves and effects of phosphorus def iciency. Plant Physiol Comm,1989,6,20-23.
[54]Yan X, Liao H, Trull MC, et al. Induction of a major leaf acid phosphatase does not confer adaptation to low phosphorus availability in common bean. Plant Physiol, 2001,125,1901-1911.
[55]张斌,秦岭.植物对低磷胁迫的适应及其分子基础.分子植物育种,2010,8(4),776-783.
[56]Nurnberger T, Abel S, Jost W, et al. Induction of an extracellular ribonuclease in cultured tomato cells upon phosphate starvation. Plant Physiol,1990,92,970-976.
[57]Loffler A, Abel S, Jost W, et al. Phosphate-regulated induction of intracellular ribonucleases in cultured tomato (Lycopersicon esculentum) cells. Plant Physiol, 1992,98,1472-1478.
[58]Goldstein AH, Baertlein DA, Danon A. Phosphate starvation stress as an experimental system for molecular analysis. Plant Mol Biol Rep,1989,7(1),7-16.
[59]Bariola PA, Howard CJ, Taylort CB, et al. The Arabidopsis ribonuclease gene RNS1 is tightly controlled in response to phosphate limitation. Plant J,1994,6(5), 673-685.
[60]Morcuende R, Bari R, Gibon Y, et al. Genome-wide reprogramming of metabolism and regulatory networks of Arabidopsis in response to phosphorus. Plant Cell Environ,2007,30,85-112.
[61]Dodds PN, Clarke AE, Newbigin E. Molecular characterization of an S-like RNase of Nicotiana alata that is induced by phosphate starvation. Plant Mol. Biol,1996, 31,227-238.
[62]Kock M, Theierl K, Stenzel I, et al. Extracellular administration of phosphate-sequestering metabolites induces ribonucleases in cultured tomato cells. Planta,1998,204,404-407.
[63]Poirier Y, Bucher M. Phosphate transport and homeostasis in Arabidopsis, In, Chris S, Elliot M, Jeff D, et al. The Arabidopsis Book, Am Soc Plant Bio,2002, 1-35.
[64]Muchhal US, Pardot JM, Raghothama KG. Phosphate transporters from the higher plant Arabidopsis thaliana. Proc. Natl. Acad. Sci. USA,1996,9(93),10519-10523.
[65]Daram P, Brunner S, Rausch C, et al. Pht2;l encodes a low-affinity phosphate transporter from Arabidopsis. Plant Cell,1999,11,2153-2166.
[66]Versaw WK, Harrison MJ. A Chloroplast phosphate transporter, PHT2;1, influences allocation of phosphate within the plant and phosphate-starvation responses. Plant Cell,2002,14,1751-1766.
[67]Rausch C, Zimmermann P, Amrhein N, et al. Expression analysis suggests novel roles for the plastidic phosphate transporter Pht2;l in auto-and heterotrophic tissues in potato and Arabidopsis. Plant J,2004,39(1),13-28.
[68]Ticconi CA, Abel S. Short on phosphate, plant surveillance and countermeasures, Trends Plant Sci,2004,9(11),548-555.
[69]Nandi SK, Pant RC, Nissen P. Multiphasic uptake of phosphate by corn roots. Plant, Cell Environ,1987,10,463-474.
[70]Furihata T, Suzuki M, Sakurai H. Kinetic characterization of two phosphate uptake systems with different affinities in suspension-cultured Catharanthus roseus protoplasts. Plant, Cell Physiol,1992,33,1151-1157.
[71]Daram P, Brunner S, Persson BL, et al. Functional analysis and cell-specific expression of a phosphate transporter from tomato. Planta,1998,206,225-233.
[72]Liu H, Trieu AT, Blaylock LA. Cloning and characterization of two phosphate transporters from Medicago truncatula roots, regulation in response to phosphate and to colonization by arbuscular mycorrhizal fungi. Mol Plant Micro Interact, 1998,11,14-22.
[73]Nussaume L, Kanno S, Javot H, et al. Phosphate import in plants, focus on the PHT1 transporters. Front Plant Sci,2011,2,1-12.
[74]Paszkowski U, Kroken S, Roux C, et al. Rice phosphate transporters include an evolutionarily divergent gene specifically activated in arbuscular mycorrhizal symbiosis. Proc Natl Acad Sci,2002,99,13324-13329.
[75]Ai PH, Sun SB, Zhao JN, et al. Two rice phosphate transporters, OsPht1;2 and OsPht1;6 have different functions and kinetic properties in uptake and translocation. Plant J,2009,57,798-809.
[76]Jia HF, Ren HY, Gu M, et al. The phosphate transporter gene OsPhtl;8 is involved in phosphate homeostasis in rice. Plant Physiol,2011,156,1164-1175.
[77]Smith FW, Cybinski DH, Rae AL. Regulations of expression of genes encoding phosphate transporters in barley roots. Dordrecht, Kluwer Academic Publishers, 1999.
[78]Rae AL, Cybinski DH, Jarmey JM, et al. Characterization of two phosphate transporters from barley, evidence for diverse function and kinetic properties among members of the Phtl family. Plant Mol Biol,2003,53,27-36.
[79]Schunmann PHD, Richardson AE, Smith FE, et al. Characterization of promoter expression patterns derived from the Phtl phosphate transporter genes of barley (Hordeum vulgare L.). JExp Bot,2004,55(399),855-865.
[80]Nagy R, Vasconcelos MJ, Zhao S, et al. Differential regulation of five Phtl phosphate transporters from maize (Zea mays L.). Plant Biol,2006,8,186-197.
[81]Leggewie G, Willmitzer L, Riesmeier J W. Two cDNAs from potato are able to complement a phosphate uptake-deficient yeast mutant, identification of phosphate transporters from higher plants. Plant Cell,1997,9,381-392.
[82]Liu C, Muchhal US, Uthappa M, et al. Tomato phosphate transporter genes are differentially regulated in plant tissues by phosphorus. Plant Physiol,1998 a,116, 91-99.
[83]Nagy R, Karandashov V, Chague V, et al. The characterization of novel mycorrhiza-specific phosphate transporters from Lycopersicon esculentum and Solanum tuberosum uncovers functional redundancy in symbiotic phosphate transport in Solanaceous species. Plant J,2005,42,236-250.
[84]Xu GH, Chague V, Melamed-Bessud C, et al. Isolation and functional characterization of LePT4, a mycorrhizal enhanced phosphate transporter in tomato, In, Li C J, Zhang F S, Doberman A. (eds.). Plant Nut Food Sec, Human Health Enviro Prot, Proc of 15th Inter Plant Nut Col, Tsinghua University Press, Beijing, China,2005,90-91.
[85]Baek SH, Chung IM, Yun SJ. Molecular cloning and characterization of a tobacco leaf cDNA encoding a phosphate transporter. Mol Cell,2001,11,1-6.
[86]Harrison MJ, Dewbre GR, Liu J. A phosphate transporter from Medicago truncatula involved in the acquisition of phosphate released by arbuscular mycorrhizal fungi. Plant Cell,2002,14,2413-2429.
[87]Javot H, Penmetsa RV, Terzaghi N, et al. A Medicago truncatula phosphate transporter indispensable for the arbuscular mycorrhizal symbiosis. Proc Natl Acad Sci,2007,104(5),1720-1725.
[88]常胜合,舒海燕,童一平,等.两个小麦磷转运蛋白基因的分离、功能鉴定和表 达研究.西北植物学报,2004,24(10),1779-1785.
[89]Davies TGE, Ying J, Xu Q, et al. Expression analysis of putative high-affinity phosphate transporters in Chinese winter wheats. Plant Cell Environ,2002,5, 1325-1339.
[90]Liu JQ, Uhde-Stone C, Li AG, et al. A phosphate transporter with enhanced expression in proteoid roots of white lupin (Lupinus albus L.). Plant Soil,2001, 237,257-266.
[91]Yang CY, Wang YX, Shen H, et al. Cloning and characterizaton of a novel root-specific putative phosphate transporter in soybean (Glycine max L.), In, Li C J, Zhang FS, Doberman A. (eds.). Plant Nut Food Sec, Human Health Enviro Prot, Proc 15th Inter Plant Nut Col, Tsinghua University Press, Beijing, China,2005, 172-173.
[92]Kai M, Masuda Y, Kikuchi Y, et al. Isolation and characterization of a cDNA from Catharanthusroseus which is highly homologous with phosphate transporter. Soil Sci Plant Nutr,1997,43,227-235.
[93]Loth-Pereda V, Orsini E, Courty PE, et al. Structure and expression profile of the phosphate Phtl transporter gene family in mycorrhizal Populustrichocarpa. Plant Physiol,2011,156,2141-2154.
[94]Mudge SR, Rae AL, Diatloff E, et al. Expression analysis suggests novel roles for members of the Phtl family of phosphate transporters in Arabidopsis. Plant J, 2002,31(3),341-353.
[95]Rausch C, Daram P, Brunner S, et al. A phosphate transporter expressed in arbuscular-containing cells in potato. Nature,2001,414,462-466.
[96]李锋,李木英,潘晓华等.不同水稻品种幼苗适应低磷胁迫的根系生理生化特性.中国水稻科学,2004,18(1),48-52.
[97]Hammond JP, Bennett MJ, Bowen HC, et al. Changes in gene expression in Arabidopsis shoots during phosphate starvation and the potential for developing smart plants. Plant Physiol,2003,132,578-596.
[98]Misson J, Raghothama KG, Jain A, et al. A genome-wide transcriptional analysis using Arabidopsis thaliana Affymetrix gene chips determined plant responses to phosphate deprivation. Proc Natl Acad Sci,2005,102,11934-11939.
[99]Wasaki J, Yonetani R, Kuroda S, et al. Transcriptomic analysis of metabolic changes by phosphorus stress in rice plant roots. Plant, Cell Environ,2003,26, 1515-1523.
[100]Wasaki J, Yonetani R, Shinano T, et al. Expression of the OsPIl gene, cloned from rice roots using cDNA microarray, rapidly responds to phosphorus status. New Phytol,2003,158,239-248.
[101]Tian J, Venkatachalam P, Liao H, et al. Molecular cloning and characterization of phosphorus starvation responsive genes in common bean (Phaseolus vulgaris L.). Planta,2007,227,151-165.
[102]Calderon-Vazquez C, Ibarra-Laclette E, Caballero-Perez J, et al. Transcript profiling of Zea mays roots reveals gene responses to phosphate deficiency at the plant-and species-specific levels. J Exp Botany,2008,59(9),2479-2497.
[103]Calderon-Vazquez C, Sawers RJH, Herrera-Estrella L. Phosphate deprivation in maize, genetics and genomics. Plant Physiol,2011,156,1067-1077.
[104]Wu P, Ma L, Hou XL, et al. Phosphate starvation triggers distinct alterations of genome expression in srabidopsis roots and leaves. Plant Physiol,2003,132, 1260-1271.
[105]Li LH, Liu C, Lian XM. Gene expression profiles in rice roots under low phosphorus stress. Plant Mol Biol,2010,72,423-432.
[106]Cai HM, Xie WB, Zhu T, et al. Transcriptome response to phosphorus starvation in rice. Acta Physiol Plant,2012,34,327-341.
[107]Gallou A, Declerck S, Cranenbrouck S. Transcriptional regulation of defence genes and involvement of the WRKY transcription factor in arbuscular mycorrhizal potato root colonization. Funct Integr Genomics,2012,12,183-198.
[108]Ren F, Chen L, Guo QQ, et al. Identification and expression analysis of genes induced by phosphate starvation in leaves and roots of Brassica napus. Plant Growth Regul,2011,65,65-81.
[109]Haran S, Logendra S, Seskar M, et al. Characterization of Arabidopsis acid phosphatase promoter and regulation of acid phosphatase expression. Plant Physiol,2000,124,615-626.
[110]Burleigh SH, Harrison MJ. The down-regulation of Mt4-like genes by phosphate fertilization occurs systemically and involves phosphate translocation to the shoots. Plant Physiol,1999,119,241-248.
[111]Liu C, Muchhal US, Raghothama KG. Differential expression of TPS11, a phosphate starvation-induced gene in tomato. Plant Mol Biol,1997,33,867-874.
[112]Martin AC, Del Pozo JC, Iglesias J, et al. Influence of cytokinins on the expression of phosphate starvation responsive genes in Arabidopsis. Plant J,2000, 24,559-567.
[113]Muchhal US, Raghotama KG. Transcriptional regulation of plant phosphate transporters. Proc Natl Acad Sci,1999,96,5868-5872.
[114]Mitsukawa N, Okumura S, Shirano Y, et al. Overexpression of an Arabidopsis thaliana high-affinity phosphate transporter gene in tobacco cultured cells enhances cell growth under phosphate-limited conditions. Proc Natl Acad Sci, 1997,94,7098-7102.
[115]Park MR, Baek SH, De los Reyes BG, et al. Overexpression of a high-affinity phosphate transporter gene from tobacco (NtPTI) enhances phosphate uptake and accumulation in transgenic rice plants. Plant Soil,2007,292,259-269.
[116]Coelho GTCP, Carneiro NP, Karthikeyan AS, et al. A phosphate transporter promoter from Arabidopsis thaliana AtPHT1;4 gene drives preferential gene expression in transgenic maize roots under phosphorus starvation. Plant Mol Biol Rep,2010,28,717-723.
[117]Shin H, Shin HS, Dewbre GR, et al. Phosphate transport in Arabidopsis, Phtl;1 and Phtl;4 play a major role in phosphate acquisition from both low-and high-phosphate environments. Plant J,2004,39,629-642.
[118]Nagarajan VK, Jain A, Poling MD, et al. Arabidopsis Phtl;5 mobilizes phosphate between source and sink organs and influences the interaction between phosphate homeostasis and ethylene signaling. Plant Physiol,2011,156,1149-1163.
[119]Liu F, Wang ZY, Ren HY, et al. OsSPX1 suppresses the function of OsPHR2 in the regulation of expression of OsPT2 and phosphate homeostasis in shoots of rice. Plant J,2010,62,508-517.
[120]Sun SB, Gu M, Cao Y, et al. A constitutive expressed phosphate transporter, OsPhtl;1, modulates phosphate uptake and translocation in phosphate-replete rice. Plant Physiol,2012,8(159),1571-1581.
[121]Baldwin JC, Karthikeyan AS, Cao AQ, et al. Biochemical and molecular analysis of LePS2;l, a phosphate starvation induced protein phosphatase gene from tomato. Planta,2008,228,273-280.
[122]Baldwin JC, Karthikeyan AS, Raghothama KG. LEPS2, a phosphorus starvation-induced novel acid phosphatase from tomato. Plant Physiol,2001,125, 728-737.
[123]Martin T, Hellmann H, Schmidt R, et al. Identification of mutants in metabolically regulated gene expression. Plant J.1997,11,53 62.
[124]Hou X L, Wu P, Jiao FC, et al. Regulation of the expression of OsIPSl and OsIPS2 in rice via systemic and local Pi signalling and hormones. Plant, Cell Environ. 2005,28,353 364.
[125]Muller R, Morant M, Jarmer H, et al. Genome-wide analysis of the Arabidopsis leaf transcriptome reveals interaction of phosphate and sugar metabolism. Plant Physiol,2001,143,156-171.
[126]Rubio V, Linhares F, Solano R, et al. A conserved MYB transcription factor involved in phosphate starvation signaling both in vascular plants and in unicellular algae. Genes Dev,2001,15,2122-2133.
[127]Nilsson L, Muller R, Nielsen TH. Increased expression of the MYB-related transcription factor, PHR1, leads to enhanced phosphate uptake in Arabidopsis thaliana. Plant, Cell Environ,2007,30,1499-1512.
[128]Devaiah BN, Karthikeyan AS, Raghothama KG. WRKY75 transcription factor is a modulator of phosphate acquisition and root development in Arabidopsis. Plant Physiol,2001,143,1789-1801.
[129]Devaiah BN, Nagarajan VK, Raghothama KG. Phosphate homeostasis and root development in Arabidopsis is synchronized by the zinc finger transcription factor ZAT6. Plant Physiol,2007,145,147-159.
[130]Devaiah BN, Madhuvanthi R, Karthikeyan AS, et al. Phosphate starvation responses and gibberellic acid biosynthesis are regulated by the MYB62 transcription factor in Arabidopsis. Mol Plant,2009,2,43-58.
[131]Chen YF, Li LQ, Xu Q, et al. The WRKY6 transcription factor modulates PHOSPHATE1 expression in response to low Pi stress in Arabidopsis. Plant Cell, 2009,21,3554-3566.
[132]Chen ZH, Nimmo GA, Jenkins GI, et al. BHLH32 modulates several biochemical and morphological processes that respond to Pi starvation in Arabidopsis. Biochem J,2007,405,191-198.
[133]Yi KK, Wu Z, Zhou J, et al. OsPTFl, a novel transcription factor involved in tolerance to phosphate starvation in rice. Plant Physiol,2005,138,2087-2096.
[134]Tang Z, Sadka A, Morishige D T, et al. Homeodomain leucine zipper proteins bind to the phosphate response domain of the soybean VspB tripartite promoter. Plant Physiol,2001,125,797-809.
[135]Wykoff DD, Grossman AR, Weeks DP, et al. Psrl, a nuclear localized protein that regulates phosphorus metabolism in Chlamydomonas. Proc Natl Acad Sci,1999, 96,15336-15341.
[136]Franco-Zorrilla JM, Gonzalez E, Bustos R, et al. The transcriptional control of plant responses to phosphate limitation. JExp Botany,2004,55(396),285-293,
[137]Schunmann PHD, Richardson AE, Vickers C E, et al. Promoter analysis of the barley Phtl;l phosphate transporter gene identifies regions controlling root expression and responsiveness to phosphate deprivation. Plant Physiol,2004,136, 4205-4214.
[138]Del Pozo JC, Allona I, Rubio V, et al. A type 5 acid phosphatase gene from Arabidopsis thaliana is induced by phosphate starvation and by some other types of phosphate mobilizing/oxidative stress conditions. Plant J,1999,19,579-589.
[139]Li D, Zhu H, Liu K, et al. Purple acid phosphatases of Arabidopsis thaliana, comparative analysis and differential regulation by phosphate deprivation. J Biol Chem,2002,277,27772-27781.
[140]Okumura S, Mitsukawa N, Shirano Y, et al. Phosphate transporter gene family of Arabidopsis thaliana. DNA Res,1998,5,261-269.
[141]Mudge SR, Rae AL, Diatloff E, et al. Expression analysis suggests novel roles for members of the Phtl family of phosphate transporters in Arabidopsis. Plant J, 2002,31,341-353.
[142]Taylor CB, Bariola PA, del Cardayre SB, et al. RNS2, a senescence-associated RNase of Arabidopsis that diverged from the S-RNases before speciation. Proc Natl Acad Sci,1993,90,5118-5122.
[143]Schachtman DP, Shin R. Nutrient sensing and signaling. Annu Rev Plant Biol,2007, 58,47-69.
[144]Miura K, Rus A, Sharkhuu A, et al. The Arabidopsis SUMO E3 ligase SIZ1 controls phosphate deficiency responses. Proc Natl Acad Sci,2005,102, 7760-7765.
[145]Bari R, Pant BD, Stitt M, et al. PHO2, microRNA399, and PHR1 define a phosphate-signaling pathway in plants. Plant Physiol,2006,141,988-999.
[146]Pant BD, Buhtz A, Kehr J, et al. MicroRNA399 is a long-distance signal for the regulation of plant phosphate homeostasis. Plant J,2008,53,731-773.
[147]Valli A, Todesco M, Mateos I, et al. Target mimicry provides a new mechanism for regulation of microRNA activity. Nat Genet,2007,39,1033-1037.
[148]Chiou TJ, Aung K, Lin SI, et al. Regulation of phosphate homeostasis by microRNA in Arabidopsis. Plant Cell,2006,18,412-421.
[149]Aung K, Lin SI, Wu CC, et al. Pho2, a phosphate overaccumulator, is caused by a nonsense mutation in a microRNA399 target gene. Plant Physiol,2006,141, 1000-1011.
[150]Fujii H, Chiou TJ, Lin SI, et al. A miRNA involved in phosphate-starvation response in Arabidopsis. Current Biol,2005,15,2038-2043.
[151]Lin SI, Chiang SF, Lin WY, et al. Regulatory network of microRNA399 and PHO2 by systemic signaling. Plant Physiol,2008,147,732-746.
[152]Franco-Zorrilla JM, Valli A, Todesco M, et al. Target mimicry provides a new mechanism for regulation of microRNA activity. Nat Genet,2007,39,1033-1037.
[153]Gonzalez E, Solano R, Rubio V, et al. Phosphate Transporter Traffic Facilitator 1 is a plant-specific SEC12-related protein that enables the endoplasmic reticulum exit of a high affinity phosphate transporter in Arabidopsis. Plant Cell.2005,17, 3500-3512.
[154]Zhou J, Jiao FC, Wu ZC, et al. OsPHR2 is involved in phosphate-starvation signaling and excessive phosphate accumulation in shoots of plants. Plant Physiol, 2008,146,1673-1686.
[155]Wu P, Wang XM. Role of OsPHR2 on phosphorus homeostasis and root hairs development in rice (Oryza sativa L.). Plant Signal Beh,2008,3,674-675.
[156]Kaeppler SM, Parke JL, Mueller SM, et al. Variation among maize inbred lines and detection of quantitative trait loci for growth at low phosphorus and responsiveness to Arbuscular mycorrhizal fungi. Crop Sci,2000,40,358-363.
[157]Da Silva AE, Gabelman W. Screening maize inbred lines for tolerance to low-P stress condition. Plant Soil,1992,146,181-187.
[158]张吉海,高世斌,杨克诚,等.玉米耐低磷种质资源的筛选与鉴定.植物遗传资源学报,2008,9(3),335-339.
[159]张丽梅,贺立源,李建生,等.玉米自交系耐低磷材料苗期筛选研究.中-国农业科学,2004,37(12),1955-1959.
[160]Vance CP. Quantitative trait loci, epigenetics, sugars, and microRNAs: Quaternaries in phosphate acquisition and use. Plant Physiol,2010,154,582 588.
[161]Century K, Lynne RT, Ratcliffe O J. Regulating the regulators, the future prospects for transcription-factor-based agricultural biotechnology products. Plant physiol, 2008,147(1),20-29.
[162]明凤,米国华,张福锁,等.水稻对低磷反应的基因型差异及其生理适应机制的初步研究.应用与环境生物学报,2000,6(2),1 38-141.
[163]华瑞,沈玉芳,李世清,等.小麦/玉米苗期磷累积量对介质供磷水平反应的差异.中国生态农业学报,2009,1 7(3),429-435.
[164]孔忠新,杨丽丽,张政值,等.小麦耐低磷基因型的筛选.麦类作物学报,201 0,30(4),591-595.
[165]丁洪,李生秀.大豆品种耐低磷和对磷肥效应的遗传差异.植物营养与肥料学报,1998,4(3),257-263.
[166]Lynch J. The role of nutrient-efficient crops in modern agriculture. J Crop Prod, 1998,1(2),241-264.
[167]Barry DAJ, Miller MH. Phosphorus nutritional requirement of maize seedlings for maximum yield. Agron J,1989,81,95-99.
[168]李绍长,龚江,王军.玉米自交系苗期耐低磷基因型的筛选.玉米科学,2003,11(3),85-89.
[169]吴平,倪俊健.应用AFLP与RFLP标记研究水稻磷吸收与利用率的数量性 状位点.植物学报,2000,42(3),229-233.
[170]龚江,李绍长,夏春兰,等.低磷胁迫下玉米自交系磷高效基因型筛选.新疆农业科学,2002,39(2),77-81.
[171]杜育梅,白昌军,田江,等.柱花草适应酸性缺磷土壤的基因型差异及可能的生理机制.华南农业大学学报,2008,29(4),6-11.
[172]章爱群,贺立源,门玉英,等.磷水平对不同耐低磷玉米基因型幼苗生长和养分吸收的影响.应用与环境生物学报,2008,14(3),347-350.
[173]Martin C, Paz-Ares J. MYB transcription factors in plants. Trends Genet,1997,13, 67-73.
[174]刘蕾,杜海,唐晓凤,等.MYB转录因子在植物抗逆胁迫中的作用及其分子机理,遗传,2008,30(10),1265-1271.
[175]Livak KJ, Schmittgen TD. Analysis of relative gene expression data using real-time quantitative PCR and the 2(-Delta Delta C(T)). Methods,2001,25,402-408.
[176]Clough SJ, Bent AF. Floral dip, a simplified method for Agrobacterium-mediated transformation of Arabidopsis thaliana. Plant J,1998,16,735 743.
[177]黄国存,朱生伟,董越梅,等.绿色荧光蛋白及其在植物研究中的应用.植物学通报,1998,15(5),24-30.
[178]Alexander M, Markus I, Dieter H. Stable transformation of sunflower(Helianthus annuus L.) using a non-meristematic regeneration protocol and green fluorescent protein as a vital marker. Trans Res,2001,10,435 444.
[179]Inger BH, Henrik BP, Mette, et al. Transformation of barley (Hordeum vulgare L.) by Agrobacterium tumefaciens infection of in vitro cultured ovules. Plant Cell Rep, 2006,25,1325 1335.
[180]Abel S, Ticconi CA, Delatorre, CA. Phosphate sensing in higher plants. Physiol. Plant,2002,115,1-8.
[181]May A, Berger S, Hertel T, et al. The Arabidopsis thaliana phosphate starvation responsive gene AtPPsPasel encodes a novel type of inorganic pyrophosphatase. Biochim Biophys Acta,2011,1810,178 185.
[182]Raghothama KG. Phosphate transport and signaling. Curr Opin Plant Biol,2000,3, 182 187.