When plants face an environmental stress such as water deficit, soil salinity, high temperature, or shade, good communication between above-and belowground organs is necessary to coordinate growth and development. Various signals including hormones, peptides, proteins, hydraulic signals, and metabolites are transported mostly through the vasculature to distant tissues. How shoots and roots synchronize their response to stress using mobile signals is an emerging field of research. We summarize recent advances on mobile signals regulating shoot stomatal movement and root development in response to highly localized environmental cues. In addition, we highlight how the vascular system is not only a conduit but is also flexible in its development in response to abiotic stress. Shoot-root communication: a long-distance relationship Tissues in higher plants are highly specialized. The shoot captures solar energy by photosynthesis and carries out reproduction, whereas the root extracts water and minerals from the soil. The coordination of these specialized functions is critical for plants to thrive. The plant vascular system, comprising xylem and phloem (see Glossary), supports the plant body while also transporting many signaling molecules from shoots to roots and vice versa. Hormones are well-studied integrators of root and shoot development. Abscisic acid (ABA) [1], auxin [2], gibberellins [3], cytokinins (CKs) [4], and jasmonic acid and its relatives [5] are known to travel through the vasculature and to act in distant tissues (Figure 1, Key figure). More recently, several small peptides, proteins, and RNA molecules have been found to be mobile in xylem or phloem and to coordinate nutrient uptake and distal stress responses [6-10] (Figure 1).Localized environmental stresses such as soil salinity or light signals require controlled longdistance transport of stress signals to elicit acclimation responses at the whole-plant level [11,12]. Understanding how mobile signals activate distal stress-responsive signaling pathways and how local and distant developmental pathways are reprogrammed is an important aspect of abiotic stress tolerance. The long-distance signaling that mediates the nutrient stress response has been recently reviewed [7]. In this review we focus on recent developments in how different mobile signals coordinate shoot stomatal movements and root development in response to salinity, water-related stresses, and light and temperature changes. Furthermore, we highlight the developmental plasticity of the plant vascular system which is the central transportation path that allows mobile signals to travel over long distances in times of stress.
Bottom-up approaches: stressed roots signal to shootsRoot-derived hydraulic signals mediate the plant shoot stress response Water limitation has a profound impact on plant growth [13]. Water absorbed from the soil moves radially to the root xylem both via the apoplastic pathway and via cell-to-cell pathways through transcellular transport and the symplastic pathway. Subs...