Stomatal (mis)behaviour

When stomata first evolved they initiated the greening of terrestrial earth, and now more than 400 million years later these simple bi-cellular valves in the leaf surface regulate global fluxes of water and carbon. Despite their importance and superficial simplicity, the behaviour of stomata remains a great challenge to understand. Different approaches to studying stomatal control have yielded rather disparate models for how stomata respond to environmental stimuli. Much of this discord arises from the diversity of mechanisms apparently involved in changing guard cell turgor and hence the aperture of the stomatal pore. On the one hand, the physical tension produced by dragging water from the soil through the xylem to the leaves directly influences leaf and guard cell turgor, while on the other hand, phytohormone levels (most importantly abscisic acid), light, photosynthesis and atmospheric gases induce active changes in guard cell turgor by triggering ionic pumping. Each stomatal control mechanism has its own champion and no model has ever successfully integrated all components. In such an environment there is great value in examining how different parts of the stomatal control network interact, particularly the competition between 'hydraulic' signals related to leaf water content and 'metabolic' signals related to ambient photosynthetic conditions.