植物水通道蛋白介导的“水-肥-气”跨膜运输功能的研究进展及展望

Research progress on biological functions of plant aquaporins in water-fertilizer-gas collaborative transportation

摘要

【意义】植物水通道蛋白（aquaporin，AQP）不仅是水分跨膜运输的关键通道，还参与尿素、NH₃、CO₂等多种溶质和气体的跨膜转运，是连接植物水分、养分与气体交换的重要分子节点。然而，AQP在水–肥–气协同运输中的系统整合机制迄今缺乏全面总结，限制了对作物资源高效利用与抗逆调控网络的深入理解。因此，系统梳理AQP的多功能分工、组织定位与动态调控机制，对揭示植物水–肥–气一体化运输的分子基础具有重要理论意义和应用价值。【进展】基于51种植物基因组中2 134个AQP家族成员的比较基因组学分析，结合非洲爪蟾卵母细胞、酵母异源表达体系及拟南芥、水稻、玉米等植物的遗传生理证据，发现AQP家族规模受全基因组多倍化与生境适应双重驱动；耐旱物种中PIP和TIP亚家族显著扩张，豆科NIP明显富集，而XIP在禾本科等支系中完全丢失。结构上，AQP四聚体中央疏水孔是CO₂的优先扩散路径，单体NPA孔道负责水和尿素等小分子转运。功能上，质膜定位的PIP协同水分吸收与光合CO₂供给，液泡膜TIP参与尿素贮存与H₂O₂解毒，NIP则介导根系尿素、NH₃及硼、硅的摄取。磷酸化修饰是动态协调的核心开关：AtOST1/AtBAK1对AtPIP2;1的磷酸化可同步增强气孔CO₂导度与水分运输效率；干旱胁迫下ABA信号关闭水通道但维持中央孔CO₂通透，实现“节水保碳”。过表达OsPIP1;3、ZmPIP2;5等基因可显著提升植株水分利用效率、氮素吸收和净光合速率。综上，植物AQP通过亚家族功能分工与磷酸化等动态调控，构成水–肥–气一体化运输网络的分子枢纽。【展望】未来研究应进一步整合冷冻电镜、分子动力学模拟和单细胞时空组学等技术，深入解析AQP底物选择性、门控机制及其在细胞和器官尺度的精细调控网络。同时，应加强利用基因编辑与合成生物学等手段对PIP、TIP、NIP等关键成员进行精准改造，以创制水–肥–气协同高效、抗逆性强的作物新种质，推动AQP功能研究从模式植物向主要作物育种应用转化。

Abstract

[Significance] Plant aquaporins (AQPs) are not only key channels for transmembrane water transport but also facilitate the transport of urea, NH₃, CO₂, and other nutrients and gases, serving as crucial molecular nodes linking water, nutrient, and gas exchange in plants. However, the systemic integration mechanisms of AQPs in coordinated water–nutrient–gas transport have not been comprehensively summarized, limiting an in-depth understanding of crop resource use efficiency and stress regulatory networks. Therefore, systematically elucidating the multifunctional specialization, tissue localization, and dynamic regulation of AQPs is of great theoretical and applied significance for revealing the molecular basis of integrated water–nutrient–gas transport in plants. [Progress] Based on a comparative genomic analysis of 2 134 AQP family members identified from 51 plant genomes, combined with evidence fromXenopus oocyte and yeast heterologous expression systems and genetic and physiological studies in Arabidopsis, rice, maize, and other plants, it has been established that AQP family size is driven by both whole-genome polyploidization and habitat adaptation. In drought-tolerant species, the PIP and TIP subfamilies are significantly expanded; NIPs are markedly enriched in legumes; and XIPs are completely lost in grasses and certain other lineages. Structurally, the central hydrophobic pore of the AQP tetramer is the preferential diffusion pathway for CO₂, whereas the monomeric NPA pore is responsible for the transport of water, urea, and other small molecules. Functionally, plasma membrane-localized PIPs coordinate water uptake and photosynthetic CO₂ supply; tonoplast-localized TIPs participate in urea storage and H₂O₂ detoxification; and NIPs mediate root uptake of urea, NH₃, boron, and silicon. Phosphorylation acts as a core switch for dynamic coordination: phosphorylation of AtPIP2;1 by AtOST1/AtBAK1 can simultaneously enhance stomatal CO₂ conductance and water transport efficiency. Under drought stress, ABA signaling closes water channels while maintaining CO₂ permeability through the central pore, thereby achieving water conservation while sustaining carbon assimilation. Overexpression of OsPIP1;3, ZmPIP2;5, and other genes significantly improves plant water use efficiency, nitrogen uptake, and net photosynthetic rate. Taken together, through subfamily functional specialization and dynamic regulation such as phosphorylation, plant AQPs constitute molecular hubs of an integrated water–nutrient–gas transport network. [Prospect] Future studies should further integrate cryo-electron microscopy, molecular dynamics simulations, and single-cell spatiotemporal omics to dissect AQP substrate selectivity, gating mechanisms, and their fine regulatory networks at cellular and organ scales. Meanwhile, precision engineering of key PIP, TIP, and NIP members through gene editing and synthetic biology should be strengthened to create crop germplasm with coordinated high water–nutrient–gas use efficiency and enhanced stress tolerance, thereby promoting the translation of AQP functional studies from model plants to breeding applications in major crops.

关键词

植物水通道蛋白 / 水分运输 / 营养吸收 / 气体交换 / 分子机制

Key words

plant aquaporin / water transport / nutrient uptake / gas exchange / molecular mechanism

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基金资助

“十四五”国家重点研发项目(202IYFD1601103)

安徽省自然科学基金面上项目(240808SMC077)