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The evolution of root architecture in plants was a prerequisite for the absorption of water and minerals from the soil, and thus a major determinant of terrestrial plant colonization. Cereals have a remarkably complex root system consisting of embryonic primary roots and post-embryonic lateral roots and shoot-borne adventitious roots. Among grass species, rice adventitious roots (also called crown roots) are developed from compressed nodes at the stem base, whereas in maize, besides crown roots, several aboveground brace roots are also formed, thus adventitious root types display species-specific diversity. Despite being the backbone for the adult root system in monocots, adventitious roots are the least studied of all the plant organs. In recent times, molecular genetics, genomics and proteomics-based approaches have been utilized to dissect the mechanism of post-embryonic meristem formation and tissue patterning. Adventitious root development is a cumulative effect of the actions and interactions of crucial genetic and hormonal regulators. In this review, we provide a comprehensive view of the key regulators involved during the different stages of adventitious root development in two important crop plants, rice and maize. We have reviewed the roles of major phytohormones, microRNAs and transcription factors and their crosstalk during adventitious root development in these cereal crops.

Shoot-borne adventitious/crown roots form a highly derived fibrous root system in grasses. The molecular mechanisms controlling their development remain largely unknown. Here, we provide a genome-wide landscape of transcriptional signatures - tightly regulated auxin response and in-depth spatio-temporal expression patterns of potential epigenetic modifiers - and transcription factors during priming and outgrowth of rice (Oryza sativa) crown root primordia. Functional analyses of rice transcription factors from WUSCHEL-RELATED HOMEOBOX and PLETHORA gene families reveal their non-redundant and species-specific roles in determining the root architecture. WOX10 and PLT1 regulate both shoot-borne crown roots and root-borne lateral roots, but PLT2 specifically controls lateral root development. PLT1 activates local auxin biosynthesis genes to promote crown root development. Interestingly, O. sativa PLT genes rescue lateral root primordia outgrowth defects of Arabidopsis plt mutants, demonstrating their conserved role in root primordia outgrowth irrespective of their developmental origin. Together, our findings unveil a molecular framework of tissue transdifferentiation during root primordia establishment, leading to the culmination of robust fibrous root architecture. This also suggests that conserved factors have evolved their transcription regulation to acquire species-specific function.

We reveal the onset and dynamic tissue-specific cytokinin signaling domains and functional importance of auxin in the auxin–cytokinin interaction domains in shaping root architecture in the economically important rice plant. Plant hormones such as auxin and cytokinin are central regulators of root organogenesis. Typical in the grass species, the root system in rice is primarily composed of post-embryonic adventitious/crown roots (ARs/CRs). Antagonistic auxin–cytokinin activities mutually balance each other to ensure proper root development. Cytokinin has been shown to inhibit crown root initiation in rice; albeit, the responsive domains remain elusive during the initiation and outgrowth of crown root primordia (CRP). Here, we show the cytokinin response domains during various stages of CRP development. RNA–RNA in situ hybridization and protein immunohistochemistry studies of the reporter gene expressed under the cytokinin responsive synthetic promoter revealed detailed spatio-temporal cytokinin signaling domains in the developing CRP. Furthermore, rice lines genetically depleted for endogenous auxin in the cytokinin responsive domains provided insight into the functional importance of auxin signaling during crown root development. Thus, our study demonstrates the onset and dynamic tissue-specific cytokinin response and functional significance of auxin–cytokinin interaction during root architecture formation in rice, a model grass species.