In plants, protein-coding mRNAs can move via the phloem vasculature to distant tissues, where they may act as non-cellautonomous signals. Emerging work has identified many phloem-mobile mRNAs, but little is known regarding RNA motifs triggering mobility, the extent of mRNA transport, and the potential of transported mRNAs to be translated into functional proteins after transport. To address these aspects, we produced reporter transcripts harboring tRNA-like structures (TLSs) that were found to be enriched in the phloem stream and in mRNAs moving over chimeric graft junctions. Phenotypic and enzymatic assays on grafted plants indicated that mRNAs harboring a distinctive TLS can move from transgenic roots into wild-type leaves and from transgenic leaves into wild-type flowers or roots; these mRNAs can also be translated into proteins after transport. In addition, we provide evidence that dicistronic mRNA:tRNA transcripts are frequently produced in Arabidopsis thaliana and are enriched in the population of graft-mobile mRNAs. Our results suggest that tRNA-derived sequences with predicted stem-bulge-stem-loop structures are sufficient to mediate mRNA transport and seem to be necessary for the mobility of a large number of endogenous transcripts that can move through graft junctions.
Plasmodesmata establish a pathway for the intercellular trafficking of viral movement proteins and endogenous non-cell-autonomous proteins, such as the two closely related meristem-maintaining KNOTTED1-like homeobox (KNOX) proteins Zea mays KNOTTED1 (KN1) and Arabidopsis thaliana SHOOTMERISTEMLESS (STM). KNOX family members are DNA binding proteins that regulate the transcriptional activity of target genes in conjunction with BEL1-like homeodomain proteins. It has been shown previously, using in vivo transport assays, that the C-terminal domain of KN1, including the homeodomain, is necessary and sufficient for cell-to-cell transport through plasmodesmata. Here, using interaction and coexpression assays, we demonstrate that the microtubule-associated and viral movement protein binding protein MPB2C from Nicotiana tabacum, and its homolog in Arabidopsis, At MPB2C, are KN1/STM binding factors. Interaction between the MPB2C proteins and KN1/STM was mapped to the KN1 homeodomain, a region not essential for heterodimerization with BEL1. Expression of MPB2C in single cells prevented KN1 cell-to-cell movement. Furthermore, in vivo trichome rescue studies established that MPB2C negatively regulates KN1 association to plasmodesmata and, consequently, cell-to-cell transport. These findings are discussed in terms of the role played by MPB2C proteins in regulating the cell-to-cell trafficking of homeodomain proteins in plants.
Plants are able to reiteratively form new organs in an environmentally adaptive manner during postembryonic development. Organ formation in plants is dependent on stem cell niches (SCNs), which are located in the so-called meristems. Meristems show a functional zonation along the apical-basal axis and the radial axis. Shoot apical meristems of higher plants are dome-like structures, which contain a central SCN that consists of an apical stem cell pool and an underlying organizing center. Organ primordia are formed in the circular peripheral zone (PZ) from stem cell descendants in which differentiation programs are activated. One mechanism to keep this radial symmetry integrated is that the existing SCN actively suppresses stem cell identity in the PZ. However, how this lateral inhibition system works at the molecular level is far from understood. Here, we show that a defect in the putative carboxypeptidase ALTERED MERISTEM PROGRAM1 (AMP1) causes the formation of extra SCNs in the presence of an intact primary shoot apical meristem, which at least partially contributes to the enhanced shoot meristem size and leaf initiation rate found in the mutant. This defect appears to be neither a specific consequence of the altered cytokinin levels in amp1 nor directly mediated by the WUSCHEL/CLAVATA feedback loop. De novo formation of supernumerary stem cell pools was further enhanced in plants mutated in both AMP1 and its paralog LIKE AMP1, indicating that they exhibit partially overlapping roles to suppress SCN respecification in the PZ.
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