• ISSN 1008-505X
  • CN 11-3996/S
董旭, 王雪, 石磊, 蔡红梅, 徐芳森, 丁广大. 植物磷转运子 PHT1 家族研究进展[J]. 植物营养与肥料学报, 2017, 23(3): 799-810. DOI: 10.11674/zwyf.16224
引用本文: 董旭, 王雪, 石磊, 蔡红梅, 徐芳森, 丁广大. 植物磷转运子 PHT1 家族研究进展[J]. 植物营养与肥料学报, 2017, 23(3): 799-810. DOI: 10.11674/zwyf.16224
DONG Xu, WANG Xue, SHI Lei, CAI Hong-mei, XU Fang-sen, DING Guang-da. Advances in plant PHT1 phosphate transporter family research[J]. Journal of Plant Nutrition and Fertilizers, 2017, 23(3): 799-810. DOI: 10.11674/zwyf.16224
Citation: DONG Xu, WANG Xue, SHI Lei, CAI Hong-mei, XU Fang-sen, DING Guang-da. Advances in plant PHT1 phosphate transporter family research[J]. Journal of Plant Nutrition and Fertilizers, 2017, 23(3): 799-810. DOI: 10.11674/zwyf.16224

植物磷转运子 PHT1 家族研究进展

Advances in plant PHT1 phosphate transporter family research

  • 摘要:
    目的磷是植物生长发育所必需的大量营养元素。植物 PHT1 磷转运蛋白家族在植物磷吸收、运转及再利用等过程中发挥了重要作用。迄今已在多种高等植物中相继分离出大量 PHT1 家族基因。本文综述了国内外关于植物 PHT1 家族的主要研究进展,详细阐述了植物 PHT1 家族的表达模式、功能及可能的调控途径。
    主要进展植物 PHT1 家族属于 MFS (major facilitator superfamily) 超家族,不同物种 PHT1 家族蛋白的结构非常保守,通常具有 12 个亲脂跨膜结构域,形成“6 螺旋–亲水大环–6 螺旋”式的结构镶嵌于质膜当中。同时,该家族具有 H2PO4/nH+ 共运子、糖转运子和 MFS 通用转运子等特征结构域和一段保守的氨基酸特征序列 GGDYPLSATIMSE。一般情况,植物 PHT1 家族基因吸收转运 1 个无机磷需要 2~4 个质子协同进入质膜,并伴随膜电位的变化。植物 PHT1 家族的磷转运特性差异较大,其动力学参数 Km 值差别较大。高等植物 PHT1 家族成员众多。在拟南芥、水稻、大豆、茄科植物及其他物种中的研究发现,PHT1 家族各成员间的时空表达模式存在差异,多数成员受低磷信号调控且主要在根部表达,少部分成员在除根以外的其他器官中表达,并行使相应的磷转运功能。已有研究表明,植物 PHT1 家族基因的转录水平受到多因素的调控,例如外界环境中的无机磷浓度,转录因子如 MYB 家族、WRKY 家族以及 ZAT6 等基因能与 PHT1 家族基因启动子区的特殊调控元件如 MYCS 元件、P1BS 元件及 W-box 元件等结合,调控基因的转录。此外,部分 PHT1 家族基因的转录水平受丛枝菌根真菌 (arbuscular mycorrhizal fungi,AMF) 的调控。除了转录水平的调控,关于植物 PHT1 家族转录后水平的调控途径同样取得了较大进展。PHF1 基因、含 SPX 结构域的蛋白家族、MicroRNA、蛋白磷酸化与去磷酸化、染色质修饰及其他等一系列调控途径均参与到 PHT1 家族基因的转录后调控及信号转导。植物激素如生长素、乙烯和细胞分裂素等也参与这一调控过程。
    建议与展望植物对磷吸收利用的分子调控机理及信号转导途径十分复杂,因此,培育磷高效利用基因型作物任重而道远。关于植物 PHT1 家族基因的研究已从模式植物向作物及其他高等植物中扩展,然而对该家族蛋白的生化及结构生物学等研究还待进一步深入。同时,对于一些基因组较复杂的多倍体物种如甘蓝型油菜、小麦、大麦及棉花等,仍有待开展进一步研究。

     

    Abstract:
    ObjectivesPhosphorus (P) is one of the essential macronutrients for plants. Plant PHT1 family proteins have vital roles in plant P absorption, mobilization and re-utilization. Numerous members of the PHT1 family have been cloned from various higher plant species. In this paper, researches in plant PHT1 phosphate transporter family were summarized, including gene expression profiles, gene function and possible regulation mechanism.
    Major advancesPlant PHT1 family belongs to a major facilitator superfamily (MFS). The protein structure of PHT1 genes is extraordinarily conserved, sharing a common topology with 12 membrane-spanning domains, which are separated into two groups of six domains by a charged hydrophilic loop, with the protein inserted in the plasma membrane. Meanwhile, PHT1 family genes have the same characterized domains as in Pi: H+ co-transporters, sugar transporters, and MFS transporters as well as a conserved amino acid sequence (GGDYPLSATIMSE). Generally, two to four protons enter the plasma membrane simultaneously while plants uptake one phosphate ion via PHT1 family genes, with changes in membrane potential. There are massive members of PHT1 family in plants which are rather different from each other in transport characteristics. The variation of PHT1 gene sequences can give rise to significant changes of their kinetic parameters of Km value. Until now, a huge number of PHT1 family members have been identified in higher plants such as arabidopsis, rice, soybean, solanaceae etc. The expression profiles of PHT1 family genes vary largely. It is generally recognized that most of the PHT1 family members can be regulated by the signal of low P stress and express in plant roots, while the expression of part of the family members can be detected in other organs of plants, and these genes function distinctly. The transcriptional levels of PHT1 family genes can be regulated by many factors such as external inorganic P concentration. Some transcriptional factors from MYB family and WRKY family as well as ZAT6 can interact with PHT1 family genes by binding with some special cis-elements existing in the promoter region of plant PHT1 family genes, such as MYCS element, P1BS element and W-box element, to regulate the expression levels of PHT1 family genes. Besides, the expression levels of PHT1 family genes can also be regulated by arbuscular mycorrhiza fugi (AMF). In addition to transcriptional regulation, post-transcriptional regulation and signal transduction pathway involved in the expression of PHT1 family genes are reported by different research groups. It shows that numerous genes including PHF1, genes containing SPX domain, MicroRNA, and other genes related to protein phosphorylation and chromatin modification are involved in the process of post-transcriptional regulation of PHT1 family members, as well as plant hormones such as auxin, ethylene and cytokinin etc.
    Suggestions and ExpectationsThere is an increasingly number of reports on the mechanism of how plants response to low P stress until now, and some genes are cloned involving P absorption and utilization. The results show that complex mechanism exists in the process of P absorption and utilization as well as signal transduction pathway. Thus, it is a long way to breeding genetically modified cultivars with enhanced P efficiency. In terms of researches on PHT1 family genes, it has changed recently from model plants to crops and other higher plants. However, further researches might be conducted on biochemistry and structural biology of proteins of PHT1 family members. In addition, more researches about PHT1 family genes might also be performed towards polyploid species such as Brassica napus, wheat, barley, cotton etc.

     

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