Abstract

Cytosolic pH homeostasis is a precondition for the normal growth and stress responses in plants, and H+ flux across the plasma membrane is essential for cytoplasmic pH control. Hence, this review focuses on seven types of proteins that possess direct H+ transport activity, namely, H+-ATPase, NHX, CHX, AMT, NRT, PHT, and KT/HAK/KUP, to summarize their plasma-membrane-located family members, the effect of corresponding gene knockout and/or overexpression on cytosolic pH, the H+ transport pathway, and their functional regulation by the extracellular/cytosolic pH. In general, H+-ATPases mediate H+ extrusion, whereas most members of other six proteins mediate H+ influx, thus contributing to cytosolic pH homeostasis by directly modulating H+ flux across the plasma membrane. The fact that some AMTs/NRTs mediate H+-coupled substrate influx, whereas other intra-family members facilitate H+-uncoupled substrate transport, demonstrates that not all plasma membrane transporters possess H+-coupled substrate transport mechanisms, and using the transport mechanism of a protein to represent the case of the entire family is not suitable. The transport activity of these proteins is regulated by extracellular and/or cytosolic pH, with different structural bases for H+ transfer among these seven types of proteins. Notably, intra-family members possess distinct pH regulatory characterization and underlying residues for H+ transfer. This review is anticipated to facilitate the understanding of the molecular basis for cytosolic pH homeostasis. Despite this progress, the strategy of their cooperation for cytosolic pH homeostasis needs further investigation.

Highlights

  • As a fundamental activity in all living cells [1], cytosolic pH homeostasis is essential for the normal growth and stress responses of plants [2,3]

  • In comparison with numerous reviews that concentrate on the organelle-located proteins which are responsible for H+ flux across the endomembrane [1,6,7], summaries regarding proteins that are directly involved in H+ efflux/influx across the plasma membrane are scarce, except regarding plasma membrane H+ -ATPases [14,17]

  • The single knockout of AHA7 (Arabidopsis H+ -ATPase 7) significantly reduces the H+ efflux capacity in the root hair zone under low-phosphorus stress [39]. All these results indicate the contribution of plasma membrane H+ -ATPases to cytosolic pH control, but the direct measurement of cytosolic pH changes upon their mutation/overexpression is still lacking

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Summary

Introduction

Publisher’s Note: MDPI stays neutral with regard to jurisdictional claims in published maps and institutional affiliations. As a fundamental activity in all living cells [1], cytosolic pH homeostasis is essential for the normal growth and stress responses of plants [2,3]. This is because basic cytosolic processes such as biochemical reactions, protein stability, ion channel/transporter activity, compartmental integrity, and membrane trafficking have strict pH requirements [1,4]. In comparison with numerous reviews that concentrate on the organelle-located proteins which are responsible for H+ flux across the endomembrane [1,6,7], summaries regarding proteins that are directly involved in H+ efflux/influx across the plasma membrane are scarce, except regarding plasma membrane H+ -ATPases [14,17]. Roles of Plasma Membrane H+ -ATPases and Multiple Transporters in Cytosolic pH Homeostasis

Plasma-Membrane-Located Family Members, Function and the Effect of Their
Mechanism of H+ Transport
Regulation by Extracellular/Cytosolic pH
CHX Family
Plasma-Membrane-Located Family Members, Function, and the Effect of Their
Not All Plasma Membrane Transporters Possess H+ -Coupled Substrate Transport
Special Caution Is Needed When Drawing Conclusion to the H+ Transfer Mechanism of Transporters
Increasing Yield
Acid Stress Resistance
Conclusions and Prospects
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