  Plants both lose water and take in carbon dioxide through microscopic
stomatal pores, each of which is regulated by a surrounding pair of guard
cells. During drought, the plant hormone abscisic acid (ABA) inhibits stomatal
opening and promotes stomatal closure, thereby promoting water conservation.
Here we synthesize experimental results into a consistent guard cell signal
transduction network for ABA-induced stomatal closure, and develop a dynamic
model of this process. Our model captures the regulation of more than forty
identified network components, and accords well with previous experimental
results at both the pathway and whole cell physiological level. Our analysis
reveals the novel predictions that the disruption of membrane depolarizability,
anion efflux, actin cytoskeleton reorganization, cytosolic pH increase, the
phosphatidic acid pathway or of K+ efflux through slowly activating K+ channels
at the plasma membrane lead to the strongest reduction in ABA responsiveness.
Initial experimental analysis assessing ABA-induced stomatal closure in the
presence of cytosolic pH clamp imposed by the weak acid butyrate is consistent
with model prediction. Our method can be readily applied to other biological
signaling networks to identify key regulatory components in systems where
quantitative information is limited.
