Guard cells perceive a variety of chemicals produced metabolically in response to abiotic and biotic stresses, integrate the signals into reactive oxygen species and calcium signatures, and convert these signatures into stomatal movements by regulating turgor pressure. Guard cell behaviors in response to such complex signals are critical for plant growth and sustenance in stressful, ever-changing environments. The key open question is how guard cells achieve the signal integration to optimize stomatal aperture. Abscisic acid is responsible for stomatal closure in plants in response to drought, and its signal transduction has been well studied. Other plant hormones and low-molecular-weight compounds function as inducers of stomatal closure and mediators of signaling in guard cells. In this review, we summarize recent advances in research on the diverse stomatal signaling pathways, with specific emphasis on signal integration and signal interaction in guard cell movement.

Methyl jasmonate (MeJA) elicits stomatal closing similar to abscisic acid (ABA), but whether the two compounds use similar or different signaling mechanisms in guard cells remains to be clarified. We investigated the effects of MeJA and ABA on second messenger production and ion channel activation in guard cells of wild-type Arabidopsis (Arabidopsis thaliana) and MeJA-insensitive coronatine-insensitive 1 (coi1) mutants. The coi1 mutation impaired MeJA-induced stomatal closing but not ABA-induced stomatal closing. MeJA as well as ABA induced production of reactive oxygen species (ROS) and nitric oxide (NO) in wild-type guard cells, whereas MeJA did not induce production of ROS and NO in coi1 guard cells. The experiments using an inhibitor and scavengers demonstrated that both ROS and NO are involved in MeJA-induced stomatal closing as well as ABA-induced stomatal closing. Not only ABA but also MeJA activated slow anion channels and Ca(2+) permeable cation channels in the plasma membrane of wild-type guard cell protoplasts. However, in coi1 guard cell protoplasts, MeJA did not elicit either slow anion currents or Ca(2+) permeable cation currents, but ABA activated both types of ion channels. Furthermore, to elucidate signaling interaction between ABA and MeJA in guard cells, we examined MeJA signaling in ABA-insensitive mutant ABA-insensitive 2 (abi2-1), whose ABA signal transduction cascade has some disruption downstream of ROS production and NO production. MeJA also did not induce stomatal closing but stimulated production of ROS and NO in abi2-1. These results suggest that MeJA triggers stomatal closing via a receptor distinct from the ABA receptor and that the coi1 mutation disrupts MeJA signaling upstream of the blanch point of ABA signaling and MeJA signaling in Arabidopsis guard cells.

Plants are often exposed to abiotic stresses such as drought, salinity, heat, cold, and heavy metals that induce complex responses, which result in reduced growth as well as crop yield. Phytohormones are well known for their regulatory role in plant growth and development, and they serve as important chemical messengers, allowing plants to function during exposure to various stresses. Seed priming is a physiological technique involving seed hydration and drying to improve metabolic processes prior to germination, thereby increasing the percentage and rate of germination and improving seedling growth and crop yield under normal and various biotic and abiotic stresses. Seed priming allows plants to obtain an enhanced capacity for rapidly and effectively combating different stresses. Thus, seed priming with phytohormones has emerged as an important tool for mitigating the effects of abiotic stress. Therefore, this review discusses the potential role of priming with phytohormones to mitigate the harmful effects of abiotic stresses, possible mechanisms for how mitigation is accomplished, and roles of priming on the enhancement of crop production.

Several recent studies have shown that citric acid/citrate (CA) can confer abiotic stress tolerance to plants. Exogenous CA application leads to improved growth and yield in crop plants under various abiotic stress conditions. Improved physiological outcomes are associated with higher photosynthetic rates, reduced reactive oxygen species, and better osmoregulation. Application of CA also induces antioxidant defense systems, promotes increased chlorophyll content, and affects secondary metabolism to limit plant growth restrictions under stress. In particular, CA has a major impact on relieving heavy metal stress by promoting precipitation, chelation, and sequestration of metal ions. This review summarizes the mechanisms that mediate CA-regulated changes in plants, primarily CA’s involvement in the control of physiological and molecular processes in plants under abiotic stress conditions. We also review genetic engineering strategies for CA-mediated abiotic stress tolerance. Finally, we propose a model to explain how CA’s position in complex metabolic networks involving the biosynthesis of phytohormones, amino acids, signaling molecules, and other secondary metabolites could explain some of its abiotic stress-ameliorating properties. This review summarizes our current understanding of CA-mediated abiotic stress tolerance and highlights areas where additional research is needed.