Membrane Transport in Guard Cells

Abstract

One of the most fascinating processes in land plants is the ability to regulate gas exchange and transpiration by the opening and closing of the stomatal pore. The guard cells of stomata serve dual, sometimes conflicting functions in regulating photosynthetic carbon dioxide uptake while minimising water loss by transpiration. Guard cells incorporate complex, flexible and robust membrane transport and control mechanisms. The dominant pathways for potassium, anion, calcium and proton transport have been characterised, both at the plasma membrane and tonoplast, much of this work drawing on electrophysiological techniques such as voltage and patch clamping. Recent genetic and genomic advances have led to the identification of the corresponding genes for several of these transporters. This article summarises the recent progress in guard cell membrane transport and outlooks for a systems biological approach using quantitative kinetic and homeostatic modelling to guide future research work.

Key Concepts:

  • Guard cells depend on concurrent regulation of parallel and antiparallel transport pathways.

  • Plasma membrane and tonoplast transport must be coordinated for stomatal function.

  • Quantitative approaches to kinetic and homeostatic modelling in guard cells will greatly benefit stomatal biology.

Keywords: stomatal regulation; ion channels; proton and calcium pumps; cotransporters; kinetic and homeostatic modelling

Figure 1.

Schematic diagrams for ion and solute transport across the plasma membrane and tonoplast during stomatal opening and closure. Symbols for channels, ATPases and transporters are colour coded as indicated (bottom). Images on the top are representative of Vicia faba guard cells in the open (a) and the closed (b) state. ABCB, ATP Binding Cassette transporter; ACA, Ca2+‐ATPase; AHA, plasma membrane H+‐ATPase; AKT, KAT and KC, K+ inward rectifying channels; AVP, vacuolar H+/K+‐PPase; CAX, Ca2+/H+ antiporter; CLC, anion channel/anion/H+ antiporter; CHX, Cation/H+ exchanger; GORK, guard cell outward‐rectifying K+ channel; NHX, Na+(K+)/H+ antiporter; NRT, nitrate transporter; STP, monosaccharide/H+ symporter; SUT, sucrose transporter; SLAC, slow anion channel; TPC, two pore cation channel; TPK, two pore K+ channel; VHA, vacuolar H+‐ATPase.

Figure 2.

ABA regulates membrane transport in guard cell. Dashed arrows indicate there are either other signalling intermediates (e.g. hydrogen peroxide and nitric oxide) or unknown components. Arrows for the second messages that have unknown or no activation/inhibition effects on the channels and pumps are not shown in the figure.

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Further Reading

Blatt MR (2000) Cellular signaling and volume control in stomatal movements in plants. Annual Review of Cell and Developmental Biology 16: 221–241.

De Angeli A, Thomine S, Franchisse JM et al. (2007) Anion channels and transporters in plant cell membranes. FEBS Letters 581: 2367–2374.

Dreyer I and Blatt MR (2009) What makes a gate? The ins and outs of Kv‐like K+ channels in plants. Trends in Plant Science 14: 383–390.

Drozdowicz YM and Rea PA (2001) Vacuolar H+‐pyrophosphatases: from evolutionary backwaters into mainstream. Trends in Plant Science 6: 206–211.

Gaxiola RA, Palmgren MG and Schumacher K (2007) Plant proton pumps. FEBS Letters 581: 2204–2214.

Outlaw WH (2003) Integration of cellular and physiological functions of guard cells. Critical Reviews in Plant Sciences 22: 503–529.

Pandey S, Zhang W and Assmann SM (2007) Roles of ion channels and transporters in guard cell signal transduction. FEBS Letters 581: 2325–2336.

Sze H, Li X and Palmgren MG (1999) Energization of plant cell membranes by H+‐pumping ATPases: regulation and biosynthesis. Plant Cell 11: 677–689.

Ward JM, Mäser P and Schroeder JI (2009) Plant ion channels: gene families, physiology, and functional genomics analyses. Annual Review of Physiology 71: 59–82.

Willmer C and Fricker M (1996) Stomata. London: Chapman & Hall.

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Chen, Zhong‐Hua, and Blatt, Michael R(Oct 2010) Membrane Transport in Guard Cells. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1002/9780470015902.a0021630]