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Messenger RNAs (mRNAs) are the templates for protein translation but can also act as non-cell-autonomous signaling molecules. Plants input endogenous and exogenous cues to mobile mRNAs and output them to local or systemic target cells and organs to support specific plant responses. Mobile mRNAs form ribonucleoprotein (RNP) complexes with proteins during transport. Components of these RNP complexes could interact with plasmodesmata (PDs), a major mediator of mRNA transport, to ensure mRNA mobility and transport selectivity. Based on advances in the last two to three years, this review summarizes mRNA transport mechanisms in local and systemic signaling from the perspective of RNP complex formation and PD transport. We also discuss the physiological roles of endogenous mRNA transport and the recently revealed roles of non-cell-autonomous mRNAs in inter-organism communication., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Fruit consumption is fundamental to a balanced diet. The contemporary challenge of maintaining a steady food supply to meet the demands of a growing population is driving the development of strategies to improve the production and nutritional quality of fruit. Plasmodesmata, the structures that mediate symplasmic transport between plant cells, play an important role in phloem unloading and distribution of sugars and signalling molecules into developing organs. Targeted modifications to the structures and functioning of plasmodesmata have the potential to improve fruit development; however, knowledge on the mechanisms underpinning plasmodesmata regulation in this context is scarce. In this review, we have compiled current knowledge on plasmodesmata and their structural characterisation during the development of fruit organs. We discuss key questions on phloem unloading, including the pathway shift from symplasmic to apoplastic that takes place during the onset of ripening as potential targets for improving fruit quality., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2021 The Authors. Published by Elsevier Ltd.. All rights reserved.)

The long-distance transport of sugars and nutrients through the phloem is essential for the proper function and growth of vascular plants. However, in addition to essential nutrients and sugars, phloem sap also contains small molecules (e.g. hormones) as well as a diverse population of macromolecules (i.e. proteins small RNAs, and mRNAs), the endogenous functions of which remain largely unknown. Understanding the cellular origins of these mobile macromolecules, their path into and out of the phloem translocation stream, and their fate at their new destination is essential for characterizing their presumptive function. Specialized plasmodesmal connections that regulate phloem entry and exit are central to all of these processes. Here, we highlight new discoveries underscoring plasmodesmal structure and function during unloading of various molecules in the sink, and discuss how these findings shape a new view for the potential function of phloem-mobile macromolecules., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

The complex form of higher plants requires continuous, balanced transport of nutrients in the phloem. The initial step of transferring sugars, amino acids, and other materials from photosynthetic cells to the conducting sieve tubes is known as phloem loading. Three phloem loading mechanisms have been described. The first involves release of sucrose into the apoplast and subsequent retrieval by the phloem. The initial release step in this process is now known to be mediated by a new class of transporters, the SWEET proteins. In the other two loading mechanisms, polymer trapping and diffusion, sucrose passes into the phloem through cytoplasmic channels, the plasmodesmata. Recent models have shed additional light on these mechanisms and their ability to sustain the growth of even the tallest trees., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Current conceptions of sucrose export largely neglect the effect of transpiration-induced water potential gradients within leaf mesophyll, even as the mix of convection and diffusion in the pre-phloem path remains uncertain. It is also generally held that the relative importance of convection and diffusion in the pre-phloem path is controlled by the ratio of their respective mass transfer coefficients. Here, we consider pre-phloem sucrose transport in the presence of adverse water potential gradients, finding that whether convection impedes or aids sucrose delivery to the phloem is independent of the permeability of the plasmodesmata to bulk flow, and depends only on assimilation rate, path-length, and the diffusivity. For most tissues subject to transpiration, convection through plasmodesmata pushes sugar away from the phloem., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Phloem unloading represents a series of cell-to-cell transport steps transferring phloem-mobile constituents from phloem to sink tissues/organs to fuel their development or resource storage. Our analysis focuses on unloading of two major phloem-mobile constituents, sugars and water. Their unloading can occur across phloem plasma membranes (apoplasmic unloading), through plasmodesmata interconnecting phloem and sink cells (symplasmic unloading) or predominately symplasmically with an intervening post-phloem apoplasmic step. In planta studies of phloem unloading encounter substantial technical challenges in accessing phloem within a meshwork of vascular/ground tissues. Thus, current understanding of phloem-unloading mechanisms largely has been deduced from indirect experimental measures or modelling. Here we highlight recent advances in understanding phloem unloading mechanisms and identify where important knowledge gaps remain., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Communication between cells is a crucial step to coordinate organ formation and tissue patterning. In plants, the intercellular transport of metabolites and signalling molecules occur symplastically through membranous structures (named plasmodesmata) that traverse the cell wall to connect the cytoplasm and endoplasmic reticulum of neighbouring cells. This review aims to highlight the importance of symplastic communication in plant development. We revisit current literature reporting the effects of changing plasmodesmata in cell morphogenesis, organ initiation and meristem maintenance and comment on recent work involving the identification of novel plasmodesmata regulators and of mobile developmental proteins and RNA molecules. New opportunities for unravelling the dynamic regulation and function of plasmodesmata are also discussed., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

Effective intercellular communication is crucial for the survival of plants. Because plant cells are encased in rigid cell walls, direct cell-to-cell exchange of cytoplasmic content is only possible through plasmodesmata (PD), membrane-lined nanotubes that connect the cytoplasm of adjacent cells. PD are highly dynamic communication channels that can undergo various structural and functional modifications. Recent findings in the field suggest that defense signaling pathways are tightly linked to the regulation of PD, and the restriction of PD-mediated cell-to-cell communication is an essential innate immune response to microbial pathogens. Moreover, several plasma membrane-bound signaling components, including receptor-like kinases that are known to have non-cell autonomous function or pathogen perception at the cell periphery, are found to also partition to PD. These findings hint at the novel role of PD as a signaling hub for both symplasmic and cross-membrane pathways., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

In plants, the intercellular transport of simple and complex molecules can occur symplastically through plasmodesmata. These are membranous channels embedded in cell walls that connect neighbouring cells. The properties of the cell walls surrounding plasmodesmata determine their transport capacity and permeability. These cell wall micro-domains are enriched in callose and have a characteristic pectin distribution. Cell wall modifications, leading to changes in plasmodesmata structure, have been reported to occur during development and in response to environmental signals. Cell wall remodelling enzymes target plasmodesmata to rapidly control intercellular communication in situ. Here we describe current knowledge on the composition of cell walls at plasmodesmata sites and on the proteins and signals that modify cell walls to regulate plasmodesmata aperture., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Membrane-lined channels called plasmodesmata (PD) connect the cytoplasts of adjacent plant cells across the cell wall, permitting intercellular movement of small molecules, proteins, and RNA. Recent genetic screens for mutants with altered PD transport identified genes suggesting that chloroplasts play crucial roles in coordinating PD transport. Complementing this discovery, studies manipulating expression of PD-localized proteins imply that changes in PD transport strongly impact chloroplast biology. Ongoing efforts to find genes that control root and stomatal development reveal the critical role of PD in enforcing tissue patterning, and newly discovered PD-localized proteins show that PD influence development, intracellular signaling, and defense against pathogens. Together, these studies demonstrate that PD function and formation are tightly integrated with plant physiology., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Mobile transcription factors play essential roles in plant development including the control of cell identity and tissue patterning, as well as organ initiation and the induction of major developmental switches. Within the past few years, the molecules and cellular structures that regulate the movement of these signals have emerged. Here we cover some of the major findings of the past two years as they relate to the intercellular movement of multiple different families of transcription factors., (Copyright © 2012. Published by Elsevier Ltd.)

Plasmodesmata are doors in the rigid cell wall. In multicellular tissues, they allow the passage of molecules needed to create physiological gradients and, by closure, symplastic boundaries, which are necessary for the fundamental processes of plant growth, development and defence. Despite this central role in plant growth our knowledge of their contribution has been hindered by difficulties in biochemical and molecular characterisation. Recent advances in proteomic, biochemical, cell biological and genetic analysis of their structure and function is showing that plasmodesmata are plastic yet highly regulated structures. They require the perception of small molecule signals (such as reactive oxygen species) to activate local changes in the cell wall that place physical constraints on the channel. This article reviews recent evidence that highlights the roles of the membrane subcomponents both as structural elements and as environments for resident signalling molecules., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Plasmodesmata (PDs), tiny channels connecting neighboring plant cells, play big roles in the transport of metabolites, viral movement, cell fate specification and development. Many recent studies are opening our eyes to the composition and formation of PDs, as well as the function and regulation of trafficking through them. Both proteomic and genetic approaches have revealed the central importance of callose in modulating PD connectivity. Moreover, many new developmental regulators, including transcription factors as well as small RNAs (sRNAs), have been found to be mobile and essential for specifying cell fate and tissue patterning., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

The identification and study of small RNAs, including microRNAs and trans-acting small interfering RNAs, have added a layer of complexity to the many pathways that regulate plant development. These molecules, which function as negative regulators of gene expression, are now known to have greatly expanded roles in a variety of developmental processes affecting all major plant structures, including meristems, leaves, roots, and inflorescences. Mutants with specific developmental phenotypes have also advanced our knowledge of the biogenesis and mode of action of these diverse small RNAs. In addition, previous models on the cell autonomy of microRNAs may have to be revised as more data accumulate supporting their long distance transport. As many of these small RNAs appear to be conserved across different species, knowledge gained from one species is expected to have general application. However, a few surprising differences in small RNA function seem to exist between monocots and dicots regarding meristem initiation and sex determination. Integrating these unique functions into the overall scheme for plant growth will give a more complete picture of how they have evolved as unique developmental systems.

Plasmodesmata remain one of the outstanding mysteries in plant biology. In providing conduits for the exchange of small and large, informational molecules they are central to the growth, development and defence of all higher plants. In the past few years, strategies have been devised for the molecular dissection of plasmodesmal composition and function, and we are beginning to see how these enigmatic structures will become to be understood.

Non-cell-autonomous signals in the form of microRNAs and transcription factors could have important developmental functions. Plasmodesmata (PD) form a cytoplasmic network throughout the plant body and provide the means of symplasmic cell-to-cell transport in plants. Homeodomain transcription factors, small RNA molecules and viral genomic information move selectively to adjacent cells via PD microchannels. Tissue-specific expression studies of non-cell-autonomous transcription factors and RNA molecules have confirmed that their intercellular transport is a highly regulated process, which depends on the tissue, developmental stage and nature of the transported macromolecule. We have known for some time that gene-silencing signals spread both locally from cell to cell and across long distances following the source to sink transition. Recent work has provided evidence that small single-stranded silencing-induced RNAs and microRNA molecules are present in the phloem transport system of different plant species. Further, recent evidence has confirmed that the transport of silencing RNA via PD is a regulated and active process, and that an amplification-relay mechanism is in place for the long-distance spread of silencing signals.

Plasmodesmata and the phloem form a cytoplasmic network that permits direct cell-cell communication in plants. This network can mediate the trafficking of selective proteins and RNAs that may have important developmental functions. Recent work has provided evidence that protein and RNA traffic across specific interfaces of this network is regulated in a distinct manner. Progress has been made in identifying potential cellular factors that confer such regulation. These advances should promote further investigations into the mechanisms and functions of protein and RNA traffic using biochemical, cellular, genetic and molecular tools.