The precise regulation of asymmetric cell division (ACD) is essential for plant organogenesis. In Arabidopsis roots, SHORT-ROOT (SHR) functions to promote periclinal division in cortex/ endodermis initials, which generate the ground tissue patterning. Although multiple downstream transcription factors and interplaying hormone pathways have been reported, the cellular mechanism that affects SHR-mediated periclinal division remains largely unclear. Here, we found that SHR can substantially elevate reactive oxygen species (ROS) levels in Arabidopsis roots by activating respiratory burst oxidase homologs (RBOHs). Among the ROS products, hydrogen peroxide (H 2 O 2) rather than superoxide (O 2 À) was shown to play a critical role in SHR-mediated periclinal division. Scavenging H 2 O 2 could markedly impair the ability of SHR to induce periclinal division. We also show that salicylic acid (SA) can promote H 2 O 2 production by repressing CAT expression, which greatly increased periclinal division in root endodermis. As a result, middle cortex was more frequently formed in the endodermis of snc1, a mutant with accumulated endogenous SA and H 2 O 2. In addition to RBOHs, SHR also activated the SA pathway, which might contribute to the elevated H 2 O 2 level induced by SHR. Thus, our data suggest a mechanism by which SHR creates the optimal micro-environment for periclinal division by maintaining ROS homeostasis in Arabidopsis roots.

Flooding tolerance is an important trait for tomato breeding. In this study, we obtained a recessive mutant exhibiting highly enhanced submergence resistance. Phenotypical analyses showed that this resistant to flooding (rf) mutant displays slightly chlorotic leaves and spontaneous initiation of adventitious roots (ARs) on stems. The mutation was mapped to the phytochromobilin synthase gene AUREA (AU), in which a single amino acid substitution from asparagine to tyrosine occurred. In addition to the classic function of AU in phytochrome and chlorophyll biogenesis in leaves, we uncovered its novel role in mediating AR formation on stems. We further observed temporal coincidence of the two phenotypes in the rf mutant: chlorosis and spontaneous AR formation and revealed that AU functions by maintaining heme homeostasis. Interestingly, our grafting results suggest that heme might play roles in AR initiation via long-distance transport from leaves to stems. Our results present genetic evidence for the involvement of the AU–heme oxygenase-1–heme pathway in AR initiation in tomato. As fruit production and yield in the rf mutant are minimally impacted, the mutation identified in this study may provide a target for biotechnological renovation of tomato germplasm in future breeding.

Root cap not only protects root meristem, but also detects and transduces the signals of environmental changes to affect root development. The symplastic communication is an important way for plants to transduce signals to coordinate the development and physiology in response to the changing enviroments. However, it is unclear how the symplastic communication between root cap cells affects root growth. Here we exploit an inducible system to specifically block the symplastic communication in the root cap. Transient blockage of plasmodesmata (PD) in differentiated collumella cells severely impairs the root development in Arabidopsis, in particular in the stem cell niche and the proximal meristem. The neighboring stem cell niche is the region that is most sensitive to the disrupted symplastic communication and responds rapidly via the alteration of auxin distribution. In the later stage, the cell division in proximal meristem is inhibited, presumably due to the reduced auxin level in the root cap. Our results reveal the essential role of the differentiated collumella cells in the root cap mediated signaling system that directs root development.

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